Full text of "Proceedings of the California Academy of Sciences, 4th series"

See other formats

PROCEEDINGS

OF THE

CALIFORNIA ACADEMY OF SCIENCES

Volume 53
2002

California Academy of Sciences
Golden Gate Park
San Francisco, California

SCIENTIFIC PUBLICATIONS DEPARTMENT
Alan E. Leviton, Editor
Gary C. Williams, Associate Editor

Katie Martin, Managing Editor
Hillary Culhane, Editorial Assistant

ISSN 0068-547X

Printed in the United States of America
by Allen Press, Inc., Lawrence, Kansas 66044

TABLE OF CONTENTS

KHURSEVICH, G.K., J.P. KOCIOLEK, AND S.A. FEDENYA: A new genus of fossil freshwater

diatoms (Bacillariophyta: Stephanodiscaceae) from the sediments of Lake Baikal............
LE Boeur, Burney J.: Status of pinnipeds on Santa Catalina Island...................... 11

BAUER, AARON M., TRIP LAMB, AND WILLIAM R. BRANCH: A revision of Pachydactylus
scutatus (Reptilia: Squamata: Gekkonidae) with the description of a new species from
De RapA RIM TRINITY Bre cs kt ea ton ar en oe ak rele a aie Be hc OA tation Shad ah teh exnene pe,

DANIEL, THOMAS F.: New and reconsidered Mexican Acanthaceae IX. Justicia............. 37

GOSLINER, TERRENCE M., AND ANGEL VALDES: Sponging off of Proifera: New species of

cryptic dorid nudibranchs (Mollusca, Nudibranchia) from the tropical Indo-Pacific ......... 51

SHEAR, WILLIAM A.: Five new chordeumatidan millipeds from China: New species of

Vieteuma (Kashmireumatidae) and Nepalella (Megalotylidae) .................-20-2005- 63

BAUER, AARON M.: Two new species of Cyrtodactylus (Squamata: Gekkonidae) from

RANDALL, JOHN E., AND JOHN E. MCCOSKER: Parapercis lata, a new species of sandperch

etemotmnes: Lingiipedmac) trom the Central Pacthic :. 2... 2200 csc awa eek eee ne tee ee 87

WARNER, RICHARD H.: Systematics of the genus Monolena (Melastomataceae) in Central

EUTIER SGN als Sp gman ine he SAM sete Sc ee Posing kU a ee Me NOU OCH cl UOTE a SON eG a 95

GRISWOLD, CHARLES E.: A revision of the African spider genus Raecius Simon, 1892

ene Ae ORO CRAIC AS Pt ee emer Ne ag he Tate i tye ie ee co hearin eo a Oe ae Li?

STEIN, DAvID L., AND NATALIA V. CHERNOVA: First records of snailfishes (Pisces: Liparidae)
from the Galapagos Islands, with descriptions of two new species, Paraliparis darwini and

MMe AT INP CENCE OSEIISUS ate tarts areas ts a ese peti ens eharcidveh bad Ge, Wlene a)ld-tane he wis aoe ey Beek 151
PENNY, NORMAN D., ED.: A guide to the lacewings (Neuroptera) of Costa Rica............ 161

LIND Ea OM VOLUN ee epee eae toes eer eae ee we ce eR ae Lae hs BAe ge Dre ok ag 459

lil

1"
a.
m1 pirwic fapge®

= Natu vin f ye ; ; = aN He a vipat

PROCEEDINGS

OF THE

CALIFORNIA ACADEMY OF SCIENCES

ere gti G © lee

HT YO

PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES

Volume 53, No. 1, pp. 1-10, 4 plates. April 5, 2002

A New Genus of Fossil Freshwater Diatoms (Bacillariophyta:

Stephanodiscaceae) from the sediments of Lake Baikal
by |
G. K. Khursevich
Institute of Geological Sciences | APR Ll 20 02
National Academy of Sciences of Belarus |
220141 Minsk, Belarus
J. P. Kociolek | ‘

Diatom Collection, California Academy of Sciences
Golden Gate Park, San Francisco, California 94118

and

S. A. Fedenya
Institute of Geological Sciences
National Academy of Sciences of Belarus
220141 Minsk, Belarus

Three new species of diatoms are described from early Pliocene sediments of Lake Baikal,
Russia. These species all possess circular valves, with areolae arranged in radial rows, with
radiating fascicules at the valve margin. One to ten central fultoportulae are bounded by 3
satellite pores. The valve mantle is divided into distinct sections separated by hyaline
strips. Marginal fultoportulae are located regularly around the mantle and are bounded
by 3 satellite pores. A single, small, sessile rimoportula is placed on the mantle about half
the distance between 2 marginal fultoportulae. The three species are distinguished by their
size, pattern of areolation and number and placement of processes. These three species are
not easily placed in any existing genus of freshwater centric diatoms, therefore a new ge-
nus, Tertiariopsis, is proposed. The new genus is compared with close allies, including
Tertiarius, Thalassiosira, Stephanodiscus and Stephanopsis.

Lake Baikal has been well documented as a biodiversity hot-spot—containing many endemic
taxa across the spectrum of biological diversity (Kozhov 1963), and the diatoms are no exception. In
the recent flora, early workers such as Dorogostaisky (1904), Skvortzow & Meyer (1928), Jasnitzky
(1936), Skvortzow (1937) and Skabitchevsky (1936, 1984) documented many new and unusual forms
in living communities of this ancient and deep lake. And more recently, new taxa continue to be de-
scribed from amongst Baikal’s extent flora (e.g., Makarova & Pomazkina 1992; Genkal &
Popovskaya 1990; Edlund et al. 1996).

The Baikal region, and the lake itself, have also been localities from which interesting taxa have
been described from the fossil record. Cheremissinova (1971, 1973), Lupikina & Khursevich (1991),
Khursevich (1994) and Likhoshway et al. (1997) described many new taxa from the Middle-Upper
Miocene sediments of the Tunkin Hollow, a depression basin located 60 km west of the southern edge
of Lake Baikal. More recently, Khursevich and colleagues (1989, 2000) have described endemic gen-
era from the Neogene sediments of the Transbaikal area and Lake Baikal. New and apparently exinct

]

2 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES
Volume 53, No. 1

taxa of the genera Stephanodiscus and Cyclotella were described from Pleistocene sedimentary strata
of Lake Baikal (Loginova & Khursevich 1986, 1990; Khursevich 1989, 2000; Likhoshway et al.
1997: Nikiteeva & Likhoshway 1994; Likhoshway 1996; Khursevich et al. 2001).

In an effort to document floristic elements and their relationship to climate change, the Baikal
Drilling Project has extracted several cores from the lake. Ina core taken in 1996, we have found some
interesting centric taxa to which we cannot assign a genus name. Herein, we describe three species as
new, and present them in a new genus, Tertiariopsis.

MATERIALS AND METHODS

In 1996 two parallel drilling cores BDP 96-1 (200 m in length) and BDP 96-2 (100 m in length)
were taken on top of the underwater Academician Ridge of Lake Baikal (53°41'48"N; 108°21'06"E,
water depth 321 m). Sediments consist of alternating fine clay and diatom ooze intervals. The core
was sampled every 2 cm to achieve a 500-year resolution. The magnetic polarity age model based on
13 magnetic reversal-event boundaries provides a robust chronology for BDP 96 sedimentary records
for the last 5 Ma (BDP members 1997; Williams et al. 1997). Preparation of permanent slides is de-
scribed in Grachev et al. (1997). Specimens were examined and photographed by oil immersion light
microscopy (Ergaval brightfield, in Minsk; Leica DMRB in San Francisco) and by scanning electron
microscopy (SEM) (JEOL JSM-35C in Minsk, Leo 1450VP in San Francisco).

SYSTEMATIC ACCOUNT
Genus Tertiariopsis Khursevich & Kociolek, gen. nov.
TYPE SPECIES. — Tertiariopsis imperseptus Khursevich, Fedenya & Kociolek

DESCRIPTION. — Valves circular, diameter 5.0-15.3 um. Valve face more or less flat, some-
times with slightly elevated center. Areolae arranged in radial rows of unequal length, sometimes
forming radiating fascicles (in two to six rows) at the margin of the valve face. Center of the valve with
an irregular pattern of areolae or structure is lacking. There is a distinct separation of the valve face
and mantle areolae. On the valve face may be a single, slightly eccentric fultoportula or several (2-10)
fultoportulae, with usually 3 (rarely 2) satellite pores. The valve mantle is divided into distinct sec-
tions (consisting of 6-10 vertical rows of small areolae) separated by hyaline strips. Marginal
fultoportulae are located regularly around the valve mantle. Externally they appear as distinct rounded
apertures at the base of hyaline strips. Internally they have 3 satellite pores and are covered by a mar-
ginal lamina. A single small sessile rimoportula is placed on the mantle approximately 1/2 the dis-
tance between 2 marginal fultoportulae. Spines may be present.

ETYMOLOGY. — Named because the species of this genus resemble those in the genus
Tertiarius.

Tertiariopsis imperseptus Khursevich, Fedenya & Kociolek, sp. nov.
Figs. 1-4, 12-17. Figure | is the holotype.

DESCRIPTION. — Valves circular, flat or slightly convex, 5.0—-14.6 tum in diameter. Areolae
15~20 in 10 um along the radius, arranged in radial rows of unequal length. There is a distinct break
between the valve face areolae and those of the mantle. Mantle areolae are smaller than those on the
valve face, and arranged in linear rows. Cribra were not observed on any specimens. One slightly ec-
centric fultoportula with three or four satellite pores occurs on the valve face. Mantle fultoportulae are
spaced regularly around the margin (4-6/10 jum). Internally, they have three satellite pores and are

KHURSEVICH, KOCIOLEK, AND FEDENYA: FOSSIL FRESHWATER DIATOMS 3

PLATE 1. Light microscopy, valve views. Figures 1-4. Tertiariopsis imperseptus Khursevich, Fedenya & Kociolek, sp. nov.
Figures 5-7. Tertiariopsis sibericus Khursevich, Fedenya & Kociolek, sp. nov. Figures 8-11. Tertiariopsis makarovae
Khursevich & Kociolek, sp. noy. Scale bar in figure 1 = 10 jm, same for all figures.

covered by a marginal lamina. External openings of mantle fultoportulae are found at the base of
hyaline strips interrupting the pattern of mantle areolae. A sessile rimoportula is located on the mantle
at the same level with marginal fultoportulae, approximately halfway between two marginal
fultoportulae. Spines were observed on the valve face/mantle junction.

HOLOTYPE. — Slide number 3415a, BDP 96-1, core 94-1 (2-3 cm), deposited in the Khursevich
Collection, Institute of Geological Sciences, National Academy of Sciences of Belarus, Minsk.

ISOTYPES. — Slide 3415b, BDP 96-1, core 94-1 (2-3 cm), deposited in the Baikal Drilling Pro-
ject Collection, Institute of Geochemistry, Siberian Branch of Russian Academy of Sciences, Irkutsk,
Russia. Type material has also been deposited in the Diatom Collection, California Academy of Sci-
ences.

TYPE LOCALITY. — Underwater Academician Ridge of Lake Baikal, Russia, borehole BDP
96-1, depth 184-194 m from sediment surface.

GEOLOGICAL ERA. — Early Pliocene.

ETYMOLOGY. — From the Latin imperseptus, to mean imperceptible, referring to its ambiguous
systematic position.

COMMENTS. — This species differs from Thalassiosira dispar (Peragallo & Heéribaud)
Serieyssol (Serieyssol et al. 1998: 178-180, figs 1-10) by the finer areolation of the valve face surface
(12-14, rarely 16 in 10 pm in T. dispar versus 15—20 in 10 pm in Tertiariopsis imperseptus), as well as

4 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES
Volume 53, No. 1

PLATE 2. SEM. Tertiariopsis imperseptus Khursevich, Fedenya & Kociolek, sp. nov. Figures 12-15. Valve exterior, show-
ing range of surface from smooth to small ridges and undulations. Fasciculated mantle is evident, as are openings of mantle
fultoportules. Spines may be few or lacking. Figures 16, 17. Internal views showing larger central areolae and smaller marginal
openings. Few central fultoportulae with three satellite pores are present. Scale bars = | ym.

KHURSEVICH, KOCIOLEK, AND FEDENYA: FOSSIL FRESHWATER DIATOMS 5

by the placement of the rimoportula (on the face-mantle border in Thalassiosira dispar, on the mantle
in Tertiariopsis imperseptus).

Tertiariopsis sibericus Khursevich, Fedenya & Kociolek, sp. nov.
Figs. 8-11, 18—23. Figure 8 is the holotype.

DESCRIPTION. — Valves circular, flat or slightly convex, 7.5—15.3 um in diameter. Areolae
12-15/10 um along a radius, arranged in radial rows of unequal length. Areolae decrease in size to-
wards the valve margin, where mantle areolae are distinctly separated from those on the valve face.
The valve mantle is divided into distinct sections separated by hyaline strips. These sections have 7-8
vertical rows of small areolae. The center of the valve has either an irregular pattern of areolae or the
center is structureless. Valve face fultoportulae, 2-10, occur in radial areolar rows near the center of
the valve and occupy the place of areolae. They have three (rarely two) satellite pores. Externally, they
may have small projections. A single sessile rimoportule is placed on the valve mantle. External ex-
pression of marginal fultoportulae (6 in 10 um) appear as distinct, rounded pores close to the edge of
the valve mantle at the base of hyaline strips. Internally they have three satellite pores.

HOLOTYPE. — Slide 3407a, BDP 96-1, core 93-1 (38-39 cm), deposited in the Khursevich Col-
lection, Institute of Geological Sciences, National Academy of Sciences of Belarus, Minsk.

IsoTYPES. — Slide 3407b, BDP 96-1, core 93-1 (38-39 cm), deposited in the Baikal Drilling
Project Collection, Institute of Geochemistry, Siberian Branch of Russian Academy of Sciences,
Irkutsk, Russia. Type material has also been deposited in the Diatom Collection, California Academy
of Sciences.

Type LOCALITY. — Underwater Academician Ridge of Lake Baikal, Russia, borehole BDP
96-1, depth 188.0—194.0 m from sediment surface.

GEOLOGICAL ERA. — Early Pliocene.

ETYMOLOGY. — Named for Siberia, the region of Russia where Lake Baikal is situated.

COMMENTS. — ‘This species differs from Thalassiosira cuitzeonensis Israde, Serieyssol & Gasse
(Serieyssol et al. 1998:180-182, figs 11-20) by the finer areolation of the valve face surface
(10-12/10 pm in T. cuitzeonensis versus 12—15/10 pm in Tertiariopsis sibericus) and the position of
the rimoportula (in Thalassiosira cuitzeonensis it is placed near one side and closest to the base of a
marginal fultoportula, whereas in Tertiariopsis sibericus it is placed approximately one-half the dis-
tance between two marginal fultoportulae).

Tertiariopsis makarovae Khursevich & Kociolek, sp. nov.
Figs. 5-7, 24-27. Figure 5 is the holotype.

DESCRIPTION. — Valves are circular with slightly convex or concave surfaces, 8.5—14.2 wm in
diameter. Areolae, up to 15 in 10 um along the radius, are arranged in radial rows of unequal length
forming radiating fascicles. The latter are separated by interfascicles, continuing to the end of the
valve mantle. The fascicles are biseriate to quadriseriate (rarely up to six rows) at the valve margin of
the valve face, becoming uniseriate towards the center. The mantle areolae are smaller in size and are
grouped in linear rows (4—5 small areolae in each row). One fultoportula with three satellite pores can
be seen near the center. Valve mantle with more or less a regular ring of marginal fultoportulae,
4-6/10 um. They have three satellite pores on the internal valve surface and open by small apertures at
the base of interfascicles externally. A single sessile rimoportula is located on the mantle halfway be-
tween two marginal fultoportulae.

HOLOTYPE. — Slide 3379a, BDP 96-1, core 92-1 (76-77 cm), deposited in the Khursevich Col-
lection, Institute of Geological Sciences, National Academy of Sciences of Belarus, Minsk.

6 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES
Volume 53, No. 1

PLATE 3. SEM. Tertiariopsis sibericus Khursevich, Fedenya & Kociolek, sp. nov. Figure 18. Valve exterior. Note numerous
openings of central fultoportulae and sectors of smaller areolae on mantle separated by hyaline strips. Figures 19-23. Internal
views. An alveolus is present, though its development is narrow to wide. Central fultoportulae have three satellite pores. Scale
bars = | pm.

KHURSEVICH, KOCIOLEK, AND FEDENYA: FOSSIL FRESHWATER DIATOMS i

PLATE 4. SEM. Tertiariopsis makarovae Khursevich & Kociolek, sp. nov. Figures 24 and 26. External views. Surface is
smooth to having elevated siliceous ridges. Groups of fine areolae are positioned on the mantle, separated by hyaline strips. Fig-
ures 25 and 27. Internal views. Central fultoportulae have 3 satellite pores. Marginal fultoportulae are prominent. A rimoportule
is positioned on the valve face:mantle junction. Scale bars = | jm.

ISOTYPE. — Slide 3379b, BDP 96-1, core 92-1 (76-77 cm), deposited in the Baikal Drilling Pro-
ject Collection, Institute of Geochemistry, Siberian Branch of Russian Academy of Sciences, Irkutsk,
Russia. Type material has also been deposited in the Diatom Collection, California Academy of Sci-
ences.

TyPE LOCALITY. — Underwater Academician Ridge of Lake Baikal, Russia, borehole BDP
96-1, depth 188.0—192.0 m from sediment surface.

GEOLOGICAL ERA. — Early Pliocene.

ETYMOLOGY. — Named in honor of our late colleague Prof. I. V. Makarova of Komarovy Botani-
cal Institute, Russian Academy of Sciences, St. Petersburg, Russia.

COMMENTS. — This species differs from 7. imperseptus by the presence of distinct areolar fasci-
cles separated by hyaline strips on the valve surface. Tertiariopsis makarovae differs from T.
sibericus in the position and number of valve face fultoportulae.

8 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES
Volume 53, No. |

DISCUSSION

The new genus Tertiariopsis differs from Tertiarius Hakansson & Khursevich (1997:21—22,
figs. 5-33) by the absence of alveoli on the valve mantle and by the different position of the
rimoportula (present near the base of costae or within a chamber of the alveolus internally in
Tertiarius versus at the same level as marginal fultoportulae on the valve mantle in Tertiariopsis).

Tertiariopsis differs from Thalassiosira Cleve (Hasle 1978; Makarova 1981, 1988; Round et al.
1990) in several important ways. First, the two genera differ in the structure of the valve face. In
Tertiariopsis areolae in uniseriate striae near the center frequently become multiseriate towards the
margin. In Thalassiosira areolae are arranged in more-or-less dense, radial, tangential or eccentric
rows. Second, the two genera differ by the structure of the valve mantle. In Tertiariopsis, the mantle is
divided into distinct sections, consisting of 6—10 vertical rows of small areolae separated by hyaline
strips. In Thalassiosira, small areolae are in vertical, diagonal or crossed rows that are not arranged
into distinct sections. Third, marginal fultoportulae position and structure differs between
Thalassiosira and Tertiariopsis. In Tertiariopsis a ring of marginal fultoportulae is placed near the
edge of the valve and covered internally by a marginal lamina. In Thalassiosira marginal fulto-
portulae are positioned at the valve face/mantle junction and lack laminae coverings. Fourth, the two
genera are distinguished by structure of rimportula. External rimportula opening lacks a tubulus in
Tertiariopsis, but a tubulus is present in Thalassiosira (except a small group of freshwater, fossil spe-
cies-see Kociolek and Khursevich 2001).

Some similarities exist between Tertiariopsis and Stephanodiscus Ehrenb. (Round 1981; Theriot
etal. 1987; Khursevich 1989; Round & Hakansson 1992), including the valve face with fasciculate ra-
dial areolae, marginal fultoportulae with 3 satellite pores around the mantle and the absence of alveoli.
The two genera differ in terms of internal cribra (domed in Stephanodiscus, apparently flat or raised
slightly in Tertiariopsis), structure of rimoportulae (external openings usually tubules in Stephano-
discus; tubules are absent in Tertiariopsis), and the relationship of marginal fultoportulae on the exter-
nal valve surface and spines (they are usually found together in Stephanodiscus, but this relationship
between spines and external fultoportulae openings is not consistent in Tertiariopsis; in fact in most
cases spines and fultoportulae openings are not positioned together in this new genus).

Tertiariopsis may also be compared with Stephanopsis Khursevich & Fedenya, another genus
described from the sediments of Lake Baikal (Khursevich et al. 2000). Both genera lack alveolar
striae, have marginal fultoportulae with 3 satellite pores, have external marginal fultoportulae open-
ings lacking tubules, and may have or lack spines. The two genera differ, however, by the presence of
thin, radial costae crossing the mantle internally in Stephanopsis and in the placement of rimoportulae
(occurring at the end of one to several costae at the margin, or near the center of the valve face in
Stephanopsis, but on the mantle exclusively in Tertiariopsis).

The genus may include French and Mexican species described by Serieyssol et al. (1998) from
fossil localities. These species, originally placed in Thalassiosira, lack alveolae and appear to have a
valve mantle dissected by hyaline strips. Transfer of these species to Tertiariopsis awaits review of
type material and further detailed SEM study.

ACKNOWLEDGMENTS

Cores for this study were made possible through the Baikal Drilling Project supported by the
National Science Foundation, the Russian Academy of Sciences, the Russian Ministry of Geology
and the Science and Technology Agency (STA) of Japan. We thank B. N. Khakhaev, D. Lykov of the
Nedra Drilling Enterprise and O. M. Khlystov of the Limnological Institute for technical assistance
with the cores, as well as A. N. Gwozdkov of the Institute of Geochemistry, I. A. Kalashnikova and
T.I. Khramtsova of the Institute of the Earth Crust for technical and logistical support. Scott Serata

KHURSEVICH, KOCIOLEK, AND FEDENYA: FOSSIL FRESHWATER DIATOMS 2)

provided technical support for the SEM at CAS. Special thanks are due to I. V. Makarova for useful
discussions on the morphology of the species belonging to this new genus.

LITERATURE CITED

BDP MemperS. 1997. Continuous paleoclimate record of the last 5 Ma. From Lake Baikal, Siberia. EOS, Ameri-
can Geophysical Union, Transactions 78:597—604.

CHEREMISSINOVA, E. A. 1971. Novye i Interesnye vidy diatomovykh vodoroslei iz neogena Pribaikalia.
Novitates Systematicae Plantarum Non-Vascularium 8:52—66.

. 1973. Diatomvaya flora neogenovykh otlozheny Pribaikalia. Novosibirsk, Nauka. 68 pp.

DOROGOSTAISKY, V. 1904. Materiaux pour servir a l’algologie du Lac Baikal et de son bassin. Bull. Soc. Imp.
Naturalistes Moscou, N.S. 18:229—265.

EDLUND, M. B., E. F. STOERMER, AND C. M. TAYLOR. 1996. Aulacoseira skvortzowii sp. nov. (Bacillariophyta),
a poorly understood diatom from Lake Baikal, Russia. Journal of Phycology 32:165-175.

GENKAL, S.I. AND G.I. PopovsKAYA. 1990. New representative of the genus Stephanodiscus Ehr. (S.
binderanus (Kiitz.) Krieg. var. baicalensis Popovsk. & Genkal var. nov.). Biology of Inland Waters: Infor-
mation Bulletin 85:27-31.

GRACHEV, M. A., YE. V. LIKHOSHWAY, AND S. S. VoroByOvA. 1997. Signals of paleoclimates of Upper Pleis-
tocene in the sediments of Lake Baikal. Russian Journal of Geology and Geophysics 38:957—980.

HAKANSSON, H. AND G. K. KHURSEVICH. 1997. Tertiarius gen. nov., a new genus in the Bacillariophyceae, the
transfer of some cyclotelloid species and comparison to closely related genera. Diatom Research 12:19—33.

HASLE, G. R. 1978. Some freshwater and brackish water species of the diatom genus Thalassiosira Cleve.
Phycologia 17:263-292.

JASNITZKY, V. 1936. Neue und interessante Diatomeen aus dem Baikal-See. Botanichesky Zhurnal 21:689—703.

KHURSEVICH, G. K. 1989. Atlas of the species of Stephanodiscus and Cyclostephanos (Bacillariophyta) from the
Upper Cenozoic sediments of USSR. F. Yu. Velichkevich, ed. Nauka i Technika, Minsk. 167 pp.

. 1994. Morphology and taxonomy of some centric diatom species from the Miocene sediments of the
Dzhilinda and Tunkin hollows. Memoirs of the California Academy of Sciences 17:269—280.

—— —., 2000. Morphological pecularities of some fossil species of Stephanodiscus (Bacillariophyta) from the
bottom sediments of Lake Baikal. Pp. 603-612 in Proceedings of the 14th International Diatom Sympo-
sium, S. Mayama, M. Idei & I. Koizumi, eds. Koeltz Scientific Books, Koenigstein.

KHURSEVICH, G. K., S.A. FEDENYA, E. B. KARABANOV, D.F. WILLIAMS, AND M.I. KuzMIN. 2000.
Stephanopsis Khursevich & Fedenya-new genus of class Centrophyceae (Bacillariophyta) from the Plio-
cene deposits of Lake Baikal. Algologia 10:106—109.

KHURSEVICH, G. K., E. B. KARABANOV, A. A. PROKOPENKO, D. F. WILLIAMS, M. I. KUZMIN, AND S. A.
FEDENYA. 2001. Biostratigraphic significance of new fossil species of diatom genera Stephanodiscus and
Cyclotella (Bacillariophyta) from Upper Cenozoic deposits of Lake Baikal, Siberia. Micropaleontology
47:1-25.

KHURSEVICH, G. K. AND J. P. KOCIOLEK. In press. New Tertiarius (Bacillariophyta: Stephanodiscaceae) species
from western North America. Proceedings of the 15th International Diatom Symposium, J. John, ed.
KHURSEVICH, G. K., A. I. MOISSEEVA, AND G. A. SUKHOVA. 1989. Concentrodiscus, anew genus of the family

Stephanodiscaceae (Bacillariophyta). Botanichesky Zhurnal 74:1660—1661.

KOCIOLEK, J. P. AND G. K. KHURSEVICH. 2001. Valve ultrastructure of new and rare fossil freshwater species of
Thalassiosira (Bacillariophyta) from China and U. S. A. Algologia 11:391—397.

Kozuov, M. M. 1963. Lake Baikal and its Life. Dr. W. Junk Publishers. The Hague. 344 pp.

LikHosHway, YE. V. 1996. Stephanodiscus khurseviczae sp. nov. from Pleistocene sediments of Lake Baikal.
Diatom Research 1|1:273-281.

LIKHOSHWAY, YE. V., G. V. POMAZKINA, AND T. A. NIKITEEVA. 1977. Centric diatoms from the Miocene de-
posits in the Baikal Rift Zone (Tunka basin). Russian Journal of Geology and Geophysics 38:1445—1 452.

LoGINova, L.P. AND G. K. KHURSEVICH. 1986. New and uncommon diatom species of the genera Cyclotella
(Kiitz.) Bréb. and Stephanodiscus Ehr. from the bottom deposits of Lake Baikal. Pp. 142-148 in New and
not enough known species of fossil animals and plants of Belarus, F. Yu. Velichrevich, ed. Nauka 1
Tekhnika, Minsk.

10 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES
Volume 53, No. 1

— — —, 1990. Fossil diatom flora of Lake Baikal. Pp. 146—176 in New representatives of fosil fauna and flora of
Belarus and other regions of the USSR, F. Yu. Velichrevich, ed. Nauka i Tekhnika, Minsk.

LUPIKINA, YE. G. AND G. K. KHURSEVICH 1991. Lobodiscus (Tscher.) Lupik. & Khurs.—a new genus in the
Class Centrophyceae. Algologia 1:67—70.

MakAROVA, I. V. 1981. Principles of the systematics of Thalassiosira Cleve and the significance of its taxo-
nomic characters. Pp. 1-14 in Proceedings of the 6th Symposium on Recent and Fossil diatoms, R. Ross, ed.
O. Koeltz, Koenigstein.

— — —. 1988. Diatom algae of the Seas of the U.S. S. R.: The genus Thalassiosira Cl. Z. I. Gleser & O. G.
Kusakin, eds. Nauka, Leningrad. 117 pp.

MakKAroVaA, I. V. AND G. V. POMAZKINA. 1992. Stephanodiscus inconspicuous sp. nov. (Bacillariophyta) from
Lake Baikal. Algologia 2:84—86.

NIKITEEVA, T. A. AND YE. U. LIKHOSHWAY. 1994. Cyclotella gracilis sp. nov. from Pleistocene materials of
Lake Baikal, Russia. Diatom Research 9:349-353.

ROUND, F. E. 1981. The diatom genus Stephanodiscus: an electron microscopic view of the classical species.
Archiv fiir Protistenkunde 124:447-465.

ROUND, F. E., R. M. CRAWFORD, AND D. G. MANN. 1990. The Diatoms. Morphology of the Genera. Cambridge
University Press, Cambridge. 747 pp.

ROUND, F. E. AND H. HAKANSSON. 1992. Cyclotelloid species from a diatomite in the Harz mountains, Ger-
many, including Pliocaenicus gen. nov. Diatom Research 7:109—125.

SERIEYSSOL, K. K., I. I. GARDUNO, AND F. GASSE. 1998. Thalassiosira dispar comb. nov. and T. cuitzeonensis
spec. nov. (Bacillariiophyceae) found in Miocene sediments from France and Mexico. Nova Hedwigia
66:177—-186.

SKABITCHEVSKY, A. P. 1936. Neue und interessante Diatomeen aus dem nordlichen Baikalsee. Botanichevsky
Zhurnal 21:705—719.

. 1984. Species Gomphonematis Ag. (Bacillariophyta) Lacus Baikal. Novitates Systematicae Plantarum
Non-Vascularium 21:51—62.

SkKvorTzow, B. W. 1937. Bottom diatoms from Olhon Gate of Baikal Lake. Philippine Journal of Science
62:293-377.

SKVORTZOW, B. W. AND C. I. MEYER. 1928. A contribution to the diatoms of Baikal Lake. Proceedings of the
Sungaree River Biological Station 1:1—55.

THERIOT, E. C., E. F. STOERMER, AND H. HAKANSSON. 1987. Taxonomic interpretation of the rimoportula of
freshwater genera in the centric diatom family Thalassiosiraceae. Diatom Research 2:251—265.

WILLIAMS, D. F., J. PECK, E. B. KARABANOV, A. A. PROKOPENKO, V. KING, AND M. I. KUZMIN. 1997. Lake
Baikal record of continental climate response to orbital insolation during the past 5 million years. Science
278:1114-1117.

PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES

‘al av —

Volume 53, No. 2, pp. 11-21, | fig., 8 tables. | ah | June 14, 2002

fome e MNCR Qigong

Status of Pinnipeds on Santa Catalina Island

by ;
Burney J. Le Boeuf
Institute of Marine Sciences, University of California
Santa Cruz, California 95064
email: leboeuf@cats.ucsc.edu

The aim of this study was to assess the number and distribution of pinnipeds on Santa
Catalina Island, the diet of California sea lions, Zalophus californianus, the incidence of
pinniped strandings, and the significance of dead pinnipeds as a source of food and
of organochlorine contamination for local bald eagles, Haliaeetus leucocephalus.
Twenty-four censuses were conducted from February to November 2000. A maximum of
70 California sea lions was recorded in March; numbers decreased to at least half this size
in summer and then increased again in the fall. The majority of sea lions were adult males
(64%), no pups were observed, and most animals inhabited the southwest side of the island,
87% of them at Seal Rocks. Market squid, Loligo opalescens, was the dominant prey in
February and March. The number of harbor seals, Phoca vitulina, peaked at 121 individu-
als during the spring breeding season, and the seals were most numerous on the southwest
side of the island. Only eight dead pinnipeds were observed during the study period, DDT
residue levels in sea lion blubber were low, and bald eagles were never observed feeding on
the carcasses. This suggests that dead pinnipeds are not an important source of food or con-
tamination for the bald eagles of Santa Catalina Island. Santa Catalina Island is a haul-out
site of minor importance for California sea lions, as reflected by low numbers observed and
the absence of breeding; colony numbers of both pinnipeds are in equilibrium.

The pinnipeds of Santa Catalina Island are of interest because of the island’s location in the
Southern California Bight (Fig. 1). First, the island is close to an ocean dumping site, and only 24 nau-
tical miles from the sewage outfall on the Palos Verdes Shelf in Los Angeles where large quantities of
DDT residues were discharged between 1948 and 1970 and still remain on the ocean floor
(MacGregor 1974; Chartrand et al. 1985). Pinnipeds, especially the California sea lion, Zalophus
californianus, accumulate high organochlorine pesticide levels from their diet (Le Boeuf and Bonnell
1971; DeLong et al. 1973; Buhler et al. 1975; Gilmartin et al. 1976; Lieberg-Clark et al. 1995).
Pinnipeds on Santa Catalina Island might exhibit greater than normal contaminant loads, compared
with animals from other islands in the Southern California Bight, if their prey are contaminated from
feeding near the dumpsites. If so, the dead bodies of these pinnipeds might be a source of contamina-
tion for scavenging birds (Garcelon 1994a). Second, due to the island’s close proximity to a large met-
ropolitan area, it supports the largest year-round resident population of humans of all the islands in
southern California, a high influx of transient tourists from spring through fall, and attendant
large-scale recreational boating activities. This might disturb the pinnipeds, limiting their number, af-
fecting their temporal and spatial use of the island, and reducing their reproduction. The aim of this pa-
per is to address the validity of this logic by presenting data on number, distribution, mortality and diet
of the pinnipeds on this island.

12 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES
Volume 53, No. 2

Reports of pinnipeds on Santa Catalina Island date back to at least the beginning of this century
(Rowley 1929). Early reports, however, were unreliable and unsystematic, failing to distinguish be-
tween California sea lions and Steller sea lions, Eumetopias jubatus. Early surveys by California Fish
and Game in the Southern California Bight focussed on sea lions but ignored the harbor seal, Phoca
vitulina. Some investigators censused the major rookeries in the Southern California Bight but omit-
ted Santa Catalina Island possibly because of its low colony numbers (e.g., Bartholomew and
Boolootian 1960; Lowry et al. 1992).

California sea lions and harbor seals are commonly found on Santa Catalina Island. California sea
lions have been recorded on this island periodically during boat and aerial censuses conducted since
the 1920s. Thirty-six censuses were conducted between 1927 and 1983 (Bonnot 1928, 1929; Fry
1939: Bureau of Marine Fisheries 1947; Bonnot and Ripley 1948; Ripley et al., 1962; Odell 1971;
Carlisle and Aplin 1966, 1971; Frey and Aplin 1970; Mate 1977; Bonnell et al. 1981; C. Oliver in
Seagars et al. 1985), most of them concentrated in the years 1975-1978, part of a large pinniped as-
sessment study of the Southern California Bight (Bonnell et al. 1981). These censuses indicate that: 1)
no breeding occurred on this island, contrary to the claim of Rowley (1929) and Bonnot (1929); 2) less
than 100 total animals were observed during the highest counts in 13 of the 18 years censused; 3) un-
usually greater numbers were observed in December 1977 (384) and March 1978 (971); 4) numbers
were always least during the summer months from April through September; 5) there has been no
growth trend in colony number, as reflected by 15 censuses in June spanning the years 1927 to 1977,
6) the majority of sea lions (94%) were observed along a 30-km segment on the southwest side of the
island between Catalina Harbor (the isthmus) and Seal Rocks, the southeastern point on the island
(Fig. 1); and 7) Seal Rocks supports the majority of sea lions (77%) found on the island.

Harbor seals in the Southern California Bight were not given as much attention as sea lions during
the first half of the century. They were first recorded on Santa Catalina Island in June 1964 (Odell
1971). Twenty-five aerial censuses were conducted during the period 1975—1987 (Bonnell et al.
1981; Stewart 1982; Hanan et al. 1986a, 1986b, 1987, 1988; Hanan 1996). Bonnell et al. (1981) noted
that harbor seals reproduce here in spring. The greatest number were recorded during March, April
and May, in the range of 120 to 205 seals with year-to-year counts being relatively stable.

The bald eagle, Haliaeetus leucocephalus, was a common resident of Santa Catalina and the
other Channel Islands from at least 1860 to the late 1950s (Kiff 1980). Although attempts to reintro-
duce them on Santa Catalina were initiated in 1980, six years after DDT production stopped in this
country, the eagles continued to exhibit high DDE residues, egg-shell thinning, and difficulty breed-
ing unassisted (Garcelon 1988; 1994a; Garcelon et al. 1989). The most likely sources of contamina-
tion are the prey they eat. Bald eagles of Santa Catalina Island feed primarily on fishes (80-86%),
birds (10-16%), mammals (1.5—3.5%) and invertebrates (0.9—2.3%) (Garcelon 1994b, 1994c). In
these studies, some bald eagles were observed scavenging on dead pinnipeds that washed up on the is-
land’s beaches. As top trophic level predators, pinnipeds accumulate high levels of this persistent
chemical from their fish and cephalopod diet (Le Boeuf and Bonnell 1971; Shaw 1971; DeLong,
Gilmartin and Simpson 1973; Anas 1974; Buhler et al. 1975; Gilmartin et al. 1976; Lieberg-Clark et
al. 1995). Thus, it is important to determine the frequency, magnitude, and importance of bald eagles
scavenging on dead pinnipeds on this island as well as the contaminant levels of the dead pinnipeds
that wash up on its shores.

The specific aims of this report are to: 1) update the status of California sea lions and harbor seals
on Santa Catalina Island by providing colony numbers by age, sex, island location, and time of year; 2)
document stranded pinnipeds by number, location, time of year, and duration of availability; 3) census
bald eagles in areas where pinnipeds are found and record the incidence of eagles feeding on dead
pinnipeds; 4) determine the DDT residue load in the blubber of dead California sea lions; and 5) esti-
mate sea lion diet from scat samples.

LE BOEUF: PINNIPEDS ON SANTA CATALINA ISLAND 13

San Miquel t. Les Angeles
Palos Verdes Shelf
Santa Barbara se
San Abooles I. |
Sarita Catalina f

West End Ean Clamente |

isthmus Cove 10 km

Catalina Maries” Halian Gardens

Lite Harber Long Pt

Pipetite Santa Catalina I. cppseyres
an Vreston Pr
Avalof Lovers Cove
China Pt
Salta Verde Pt

Seal Racks

FIGuRE |. Schematic map of Santa Catalina Island in relation to the mainland, the Palos Verdes Shelf, Los Angeles, and other
islands in the Southern California Bight.

METHODS

Twenty-four censuses were conducted on Santa Catalina Island from February to November
2000, at least one per month. Most censuses began at dawn and were conducted from a small boat mo-
toring slowly along the shoreline, supplemented during inclement weather in winter by land counts
from areas such as Little Harbor and Ben Weston Point (Fig. 1). The island was surveyed on March 10,
by circumnavigating the island to determine the areas where pinnipeds were located. Ten monthly
censuses covered the southwest side, from Avalon Bay to Catalina Harbor, where virtually all of the
pinnipeds associated with Santa Catalina Island were found. Thirteen censuses were restricted to the
principal haul-out site of California sea lions, the Seal Rocks area and vicinity. Approximately 80
hours were spent in censusing and observing activities.

Sea lions were categorized as adult males, females or immature males, juveniles, or pups (Peter-
son and Bartholomew 1967). Harbor seals were categorized as adults or juvenile/pups.

Nineteen sea lion scats were collected during February and March at Seal Rocks where sea lions
rest. Scats were processed and prey hard parts identified at Moss Landing Marine Laboratories (Moss
Landing, California) using the method described in Wiese (1999).

The location, sex, standard length, estimated age, and condition of dead pinnipeds found on
beaches and in adjacent waters was recorded. When possible, the number of days dead pinnipeds re-
mained in the area was recorded. A blubber tissue sample, from outer skin to muscle, approximately
150 grams, was excised from the medial ventrum region of six dead sea lions at the level of the axilla.
The samples were deposited in clean jars and refrigerated until organochlorine analysis was con-
ducted following the methods described by Tanabe et al. (1994) and Watanabe et al. (1999). Total
DDT residues (),»DDT) in blubber are reported in g/g wet weight and p1g/g lipid weight. );DDT refers
to all constituents of DDT and its metabolities: p, p’-DDT (1,1, 1-trichloro-2, 2-bis(p-chloropheny1)

14 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES
Volume 53, No. 2

ethane), p, p’-DDD (1, 1-dchloro-2, 2-bis (pchorophenyll ethane), and p, p’-DDE (1,1-dichloro-2,
2-bis (p-chloropheny1)) ethylene).

Bald eagles were counted along the pinniped census route and their presence and behavior near
dead pinnipeds was noted. Foraging attempts by bald eagles on fish, birds or dead pinnipeds were re-
corded.

RESULTS

California sea lions. The greatest number of sea lions recorded was 70 animals in March; counts
decreased to half this number by May (35), reached a low of less than 10 animals in June and July, and
then began to increase in the fall to the high counts previously observed in winter (Table 1). The age
and sex composition of animals that were categorized was 64% adult males, 26% females or subadult
males, and 10% juveniles. No pups were observed.

On each census, the majority of sea lions were located at Seal Rocks, near the southwestern tip of
the island, 2.5 nautical miles from Avalon and Avalon Bay. That is, a mean of 86.7 + 10.8 percent of
the sea lions counted along the entire resident area (Table 1) was found at Seal Rocks. The greatest
count of all censuses conducted specifically at Seal Rocks, was 70 sea lions on 6 March (Table 2). As-
suming that this number was 86.7% of the total sea lions present on the island, the estimated total was
81 sea lions present on the island on this date. The age and sex composition of sea lions at Seal Rocks
was similar to that of the entire southwest side of the island (Table 1).

Sea lion diet. Scat analysis revealed that two cephalopods, Market squid, Loligo opalescens, and
Octopus, Octopus spp., and three fishes, Pacific Whiting, Merluccius productus, Northern anchovy,
Engraulis mordax, and rockfish, Sebastes spp., were the principal prey consumed in February and
March (Table 3). Market squid was the most common prey, appearing in 74% of the samples. Liquid
scats, commonly observed when sea lions feed on squid, were prevalent on Seal Rocks in late March
in the location where sea lions rested. Moreover, squid was being fished in the area at the time.

Harbor seals. The greatest number of harbor seals was counted in March through May; counts
after June were a quarter to a third lower than the high count in March (Table 4). The seals were con-
centrated in large groups in March through May and pupping was observed in April. Most seals at this
time of year were observed between China Point and Ben Weston Point. For example, 92% of the seals

TABLE |. California sea lion censuses of Santa Catalina Island in 2000. All censuses covered the principal
haul-out sites of sea lions, from Seal Rocks to Catalina Harbor. The March 10 census covered the entire island.

Adult males Females and/or Juveniles Pups Unidentified Total

Date immature males
4 March 4] 17 6 0 | 65
10 March 32 8 4 0 26 70
23 March 20 2) 4 0 0 36
13 April ay 0 ] 0 0 53
12 May 18 8 9 0 0 35
15 June 3 l I 0 0 5
11 July 3 6 0 0 0 9
17 August 0 I 2 0 0 15
14 September d 5 0 0 I 13
18 October 9 9 I 0 3 22
10 0 4 0 0 14

15 November
Relative % of
identified 63.7 25.8 10.5

LE BOEUF: PINNIPEDS ON SANTA CATALINA ISLAND 15

TABLE 2. Censuses of California sea lions at Seal Rocks,Santa Catalina Island in 2000.

Adult males Females and/or Juveniles Pups Unidentified Totals
Date immature males
28 February — - _ — 41 4]
4 March 1 7 6 0 0 64
6 March 51 15 4 0 0 70
7 March 26 5 0 0 0 31
9 March 7) 9 0 0 0 30
10 March 32 8 4 0 0) 44
11 March 26 13 6 0 0 45
12 March 22. 12 8 0 0 42
13 March 18 12 6 0 0 36
19 March 26 1D 4 0 0 42
20 March 12 16 3 0 0 31
21 March 2 16 4 0 0 32
22 March 21 14 5 0 0 40
23 March 20 10 4 0 0 34
24 March — — - - 26 26
25 March - — - - 3] Si
13 April 50 0 | 0 0 51
12 May 18 6 i] 0 0 31
15 June 3 1 0 0 0 4
11 July 2 6 0 0 0 8
17 August 0 13 0 0 0 13
14 September 6 5 0 0 0 11
18 October 9 9 l 0 0 19
15 November 10 — 4 0 0 14
Relative % of
identified 59.2 30.5 10.3

counted on 13 April and 88% of the seals counted on 12 May were in this area. The largest aggrega-
tions were centered in the China Point area. During the rest of the year, harbor seals were distributed in
small numbers from Seal Rocks to Ben Weston Point; a group of 8—10 seals was observed on the east
side of the island | km west of Isthmus Cove on 10 March and 12 May. Unlike sea lions, Seal Rocks
was not a preferred site, however, 2—5 seals were found often at this site.

TABLE 3. Minimum number of individuals (MNI) and percentage frequency occurrence (% FO)
of prey species (identified from otoliths and beaks) in nineteen California sea lion fecal samples col-
lected on Santa Catalina Island, California, February and March 2000.

Prey Species

Common name Scientific name MNI % FO
Market Squid Loligo opalescens 197 13a
Pacific Whiting Merluccius productus 34 50
Northern Anchovy Engraulis mordax 5 15.8
Rockfish Sebastes spp. 4 15.8
Octopus Octopus spp 3 10.5
Unidentified — 1 53

16 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES
Volume 53, No. 2

TABLE 4. Harbor seal censuses on Santa Catalina Island in 2000. All censuses covered the
principal haul-out sites of sea lions, from Seal Rocks to Catalina Harbor. The March 10 census
covered the entire island.

Date Adults Juveniles and pups. _—_- Unidentified Total
4 March — - 8 8
10 March _ = jp 121
23 March - — a 71
13 April 57 14 - 71
12 May 2 - 80 82
15 June 5 - 19 24
11 July i ~ 14 45
17 August 5 = 5 10
14 September ~ = 5 5
18 October Def - 0 27
15 November 5) ~ 0 5

Relative % of
identified 90.4 9.6

Dead pinnipeds. During 4 of the 24 censuses, we saw seven dead pinnipeds in the water or
beaches (Table 5); an eighth carcass washed up near Isthmus Cove on the north side of the island and
was reported to us by G. Long. All stranded animals were California sea lions, except one, which was
unidentified. Four of the sea lions were found in Little Harbor. The three carcasses observed on 9 and
20 March, which were marked, remained on the beach for only one day. Evidently, they moved out to
sea with the tidal flow and were not observed again in the 4—5 days that followed. Three other car-
casses were found floating in the water; two carcasses had puncture holes indicating gulls had fed on
them; a western gull, Larus Occiodentalis, was perched on one. Gulls also were feeding on the un-
identifiable carcass on the beach at China Point.

Mean DDT in the blubber of six sea lions sampled (Table 6) was 13.0 + 12.0 g/g wet weight
(geometric mean = 9.1 ug/g wet weight) and 55.0 + 62.8 ug/g lipid weight (geometric mean = 37.0
ug/g lipid weight). Discarding the sample with the very low lipid value (ZC051300) lowers the mean
DDT value from 55 to 30 g/g lipid weight.

Bald eagle sightings. During 10 censuses totaling 80 hours in duration, we observed 15 bald ea-
gles. The eagles were observed for a total of 10 hours. Nine sightings were at Seal Rocks; three were
sightings of a pair of eagles and the rest were of single eagles. In all but one case, the eagles were
perched on the rocks above Seal Rocks or on rocks over the adjacent beach to the northwest. In addi-
tion, a pair of eagles was observed near Catalina Harbor and lone eagles were observed between Seal
Rocks and Salta Verde Point. We never saw eagles feeding on fish, birds or pinniped carcasses.

On 12 and 13 July, while not censusing pinnipeds, we observed a lone eagle feeding on fish at
Lover’s Cove, near Avalon, and two adults and a juvenile feeding on fish at Italian Gardens and Long
Point on the east side of the island.

DISCUSSION

Our observations confirm the conclusions from earlier researchers. Sea lions no longer breed on
Santa Catalina Island, as they did before the 1920s (Bonnot 1929; Rowley 1929), but rather use it ex-
clusively as a haul-out site. Fewer than 100 animals are seen usually on the island, with the greatest
number occurring in winter and the fewest in summer. The majority of sea lions reside in a 30-km area
on the southwest side of the island between Catalina Harbor and Seal Rocks. Because approximately

LE BOEUF: PINNIPEDS ON SANTA CATALINA ISLAND 17

TABLE 5. Dead pinnipeds observed during pinniped censuses at Catalina Island in 2000.

Condition

Location

Genus Sex Age

Identification

Date

lower half only
decomoposed
missing head

Little Harbor, on beach

juvenile

Zalophus
Zalophus

ZC030900-1

9 March

Little Harbor, on beach

subadult
juvenile

male
male

ZC030900-2
ZC032100

Little Harbor, on beach

Zalophus

21 March

not approachable

China Point, on beach

Unidentified
ZC041300

13 April

decomposed, fed on by gulls

fresh dead

Little Harbor, in water

adult

male

Zalophus

Isthmus Cove, on beach

adult

ZC051300 Zalophus

13 May
15 June

decomposed, fed on by gulls

1 km W of Salta Verde Pt.,

adult

male

Zalophus

ZC061500-1

in water
Salta Verde Pt., in water

decomposed

adult

male

Zalophus

ZC061500-2

86% of the sea lions on the island were counted on Seal
Rocks, one can make a reasonable prediction of total island
number from this location alone, when other parts of the is-
land are inaccessible due to inclement weather. The majority
of sea lions that use the island are adults and subadult males;
the rest are juveniles and possibly some non-parous females.

A comparison ofall available sea lion counts from Santa
Catalina Island, most of which were in June (Table 7), indi-
cates that colony size is not increasing but rather has re-
mained relatively stable during the last 80 years, even during
the years 1948 to 1970 when large quantities of DDT resi-
dues were being discharged on the nearby Palos Verdes
Shelf. Unusual peaks in number, however, were recorded,
such as 233 in June 1958 (Ripley et al. 1962), 277 in March
1977, 282 in October 1977, 384 in December 1977 and 971
in March 1978 (Bonnell et al. 1981). These high counts ap-
pear to have been associated with large concentrations of
market squid spawning near the island. Local squid fisher-
men report that squid were unusually abundant in 1977 and
1978. Our scat sample analysis confirms that sea lion diet in
late winter and early spring of 2000 was mainly market
squid. An earlier study indicated that market squid also was
the most common prey of sea lions in the fall of 1980 and
1983 (Lowry and Folk 1987). Thus, sea lions use Santa
Catalina Island as a resting place from which they make for-
aging forays in the nearby waters. When squid are abundant,
sea lion numbers may double or even increase tenfold.

The decline in sea lion numbers during the summer, a
reliable pattern over the years, may simply reflect adults de-
parting to breed on nearby rookeries, such as Santa Barbara,
San Nicolas, San Clemente, or San Miguel Islands. Breeding
on Santa Catalina Island may be discouraged by the large in-
flux of tourists and recreational boating activity during sum-
mer, especially because females require undisturbed sites to
give birth and nurse their pups. Moreover, tourists congre-
gate on the expansive sandy beaches, the preferred habitat of
breeding sea lions.

It is unlikely that dead sea lions in the vicinity are an im-
portant source of food, and hence, organochlorine contami-
nants, for the bald eagles of Santa Catalina Island. Few
pinniped carcasses, most of them sea lions, were observed
during censusing, which follows from the low number of an-
imals in residence, and also indicates that few carcasses orig-
inating elsewhere in the Southern California Bight strand
here. Three of the five carcasses observed on beaches were
present for only one day. No carcass was found near Seal
Rocks where sea lions were most numerous. No eagle was
ever seen feeding ona pinniped carcass. Moreover, the mean
DDT levels in the blubber of the dead sea lions sampled

18 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES
Volume 53, No. 2

TABLE 6. Total DDT residues in the blubber of dead California sea lions collected at Santa Catalina
Island in 2000.

_DDT Residues

Identification Age Category % Lipid Wet wt. ug/g Lipid wt. ug/g
ZC030900-1 juvenile 41.1 U2 IES:
ZC030900-22 subadult 21.9 4.0 18.3
ZC032100 juvenile 37/7) 7.6 20.2
ZC041300 adult 64.0 32. Il 50.0
ZC051300 adult 1.8 33} 150.0
ZC061500-1 adult 54.0 23.8 44.0
Arithmetic mean 36.8 13.0 55.0
Standard deviation 22.4 12.0 62.8
Geometric mean 24.4 9.1 37.0

was only 13 parts per million (ppm) wet weight, almost two orders of magnitude below the mean level
of 911 ppm wet weight observed in 1970 (Le Boeuf and Bonnell 1971) and similar to levels reported
from central California in 1988-1992 (Lieberg-Clark et al. 1995). This level is only marginally
greater than the mean level of DDE reported in Western gulls, Larus occidentalis, from Santa Catalina
Island in 1992, 7.4 ppm (range = 0.9-28.2 ppm) (Garcelon 1994c). In lieu of these points—and be-
cause Santa Catalina eagles feed primarily on fish, followed by piscivorous birds such as gulls, and
rarely on dead mammals and invertebrates (0.9—2.3%) (Garcelon 1994b, 1994c)—1t is unlikely that
the major source of the contaminant loads of bald eagles comes from stranded pinnipeds.

Santa Catalina Island supports a resident breeding population of approximately 100 harbor seals.
The majority of them inhabit an isolated, rugged section of the southwestern coast on the weather side
of the island that is not easily approached by land or sea. Their number is highest during the spring
breeding season and decreases by about half this number during the rest of the year. Census counts
vary considerably with the number of seals in the water foraging, the latter being correlated with tidal
conditions and sea state. Overall, their numbers have been relatively stable over the years as reflected
by the four censuses conducted in March and the ten censuses conducted in June (Table 8).

In conclusion, the California sea lion and harbor seal colonies on Santa Catalina are in equilib-
rium. In most years, Santa Catalina is a minor haul-out site between foraging bouts for less than 100
sea lions. As reflected by the low numbers observed, and the absence of breeding, the island is of little
importance for the California sea lion population, which is estimated at over 200,000 for the U. S.
(Forney et al. 2000). The major source of high organochlorine contaminant loads in local bald eagles
does not appear to be dead sea lions or seals because few are available, they are not available for long,
they may not be consumed frequently by eagles, and the contaminant loads of sea lions have decreased
considerably during the last three decades.

ACKNOWLEDGMENTS

I thank Joel Burt for assistance censusing and for making his boat, “Daymaker,” available; Frank
Starkey of the Catalina Island Conservancy for facilitating this work and assistance with logistics;
Steve Hansen and John Giesy for expert advice and assistance with organochlorine analysis; the
Catalina Island Conservancy for permission to conduct this research; Scott Davis and Gina Long for
assistance in the field; and James Lecky and Joe Cordaro of the National Marine Fisheries Service,
Southwest Region, for authorizing tissue and scat collection.

LE BOEUF: PINNIPEDS ON SANTA CATALINA ISLAND

‘(8861 “L86I

‘9861 “PO861) Te 19 UEURH “(CR6T) ‘Te 19 SIBBRAg UI JOATIO “D “(q “VPRGT) WYO, puL EMIS “(7R6T) HEMAIS “(186T) ‘TP 39 [ouUog “(1 61) [PO “BreP Joy saoinOg

€ 07 Jaquis090q

IOQUISAON

NG I 0 1940190
¢ 61 0 Jaquiaydag
Ol SI 0 jsnsny
SP 0 0 0 Ang
v7 LE ev pli 78 07 SZ 0 LOI oun
CL S07 cOl SI Avy
IL e81 judy
IZI L7I 611 OSI yore
Aren.iqof

$9 IZ 0 Arenuer

0007 L861 9861 S861 p86 €861 7861 8L6l LL6I 9L61 SL6l b961 yuo

Tea X

(S861) ‘Te 19 suBBeag U7 JOATIO “D pur (861) ‘Te 19 [ouUog “(LL61) WW “(OL61) Udy pue Jory *“(1L61 “9961) Udy

pur a1s]18D (161) HPO (Z961 ‘LP6T) ‘Te 30 Aotdry “(gp61) Aojdry pur youuog ‘(,p6]) souaysty out Jo neaing (6€61) Atd (6761 “876 1) IOUUO :€61-LZG] 40} SadINOg

p8e O€ Jaquiaseq
TIQuIsAON
(6 787 0 Jaqo19O
€l £6 St Jaquuiaydag
SI €€ LI isnony
6 0 9 0 Ayn
¢ 901 rl 0 6£ LOI Se 6 O€ CCT 07 bol SI Or SI une
Se LS (é Avy
€S 901 [udy
69 1L6 like Ol O€ yore
Areniqo.q
p8I LI 97 Arenue
QUO eSGl= GSUOl Um LZOL | 296i SGL6l. OL61 | G96l S96 sp9Gl. “196 — 8561 ““/e6- 9PGl — GE61”~ SEG) <S26le 276 yuo]
Ivo

‘rea JO yJuour Aq ‘/ 76 BOUTS puR]S] BUITEJeD eJURS UO Pasnsudd sUOT] vas RIWIOJ][RD [BIOL “L ATAVL

20 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES
Volume 53, No. 2

LITERATURE CITED

Anas, R. E. 1974. DDT plus PCB’s in blubber of harbor seals. Pesticides Monitoring Journal 8:12—14.

BARTHOLOMEW, G. A. ANDR. A. BOOLOOTIAN. 1960. Numbers and population structure of the pinnipeds on the
California Channel Islands. Journal of Mammalogy 441:36—375.

BONNELL, M. L., B. J. LE BOEUF, M. O. PIERSON, D. DETTMAN, AND G. D. FARRENS. 1981. Marine Mammal and
Seabird Surveys of the Southern California Bight Area, 1975-1978. Pinnipeds of the Southern California
Bight. University of California, Santa Cruz, California. Vol. III. Part 1. 535 pp. (NTIS PB81248171).

BONNOT, P. 1928. The sea lions of California. California Fish and Game 14:1—16.

. 1929. Report on the seals and sea lions of California. California Division of Fish and Game, Fisheries
Bulletin 14:1—62.

Bonnot, P. AND E. RIPLEY. 1948. The California sea lion census for 1947. California Fish and Game 34:89—92.

BUHLER, D. R., R. R. CLAEYS, AND B. R. MATE. 1975. Heavy metal and chlorinated hydrocarbon residues in
California sea lions (Zalophus californianus). Journal of the Fisheries Research Board of Canada
32:2391—2397.

BUREAU OF MARINE FISHERIES. 1947. Calfornia sea lion census for 1946. California Fish and Game 33:19—22.

CARLISLE, J. G. JR. AND J. A. APLIN. 1966. Sea lion census for 1965 including counts of other California Islands.
California Fish and Game 52:119—120.

. 1971. Sea lion census for 1970, including counts of other California pinnipeds. California Fish and
Game 57:124—127.

CHARTRAND, A. B.,S. Moy, A. N. SAFFORD, T. YOSHIMURA, AND L. A. SCHINAZI. 1985. Ocean dumping under
Los Angeles Regional Water Quality Board permit: A review of past practices, potential adverse impacts,
and recommendations for future action. California Regulations Water Quality Control Board, Los Angeles
Region, March 1985. 47 pp.

DELONG, R. L., W. G. GILMARTIN, AND J. G. SIMPSON. 1973. Premature births in California sea lions: associa-
tion with high organochlorine pollutant residue levels. Science 181:1168—1170.

Forney, K. A., J. BARLOW, M. M. Muto, M. Lowry, J. BAKER, G. CAMERON, J. MOBLEY, C. STINCHCOMB,
AND J. V. CARRETTA. 2000. U.S. Pacific Marine Mammal Stock Assessments: 2000.
NOAA-TM-NMES-SWESC-300.

Frey, H.W. AND J. A. APLIN. 1970. Sea lion census for 1969, including counts of other California pinnipeds.
California Fish and Game 56:130—133.

Fry, D. H. 1939. A winter influx of sea lions from lower California. California Fish and Game 35:245—250.

GARCELON, D. K. 1989. The reintroduction of bald eagles on Santa Catalina Island, California. M. S. thesis,
Humboldt State University, Arcata, California. 58 pp.

. 1994a. Effects of organochlorine contaminants on Bald Eagle reproduction at Santa Catalina Island. A

report to the Damage Assessment Office, U. S. Fish and Wildlife Service, Sacramento, California. 16 pp.

. 1994b. Food habits of the bald eagle on Santa Catalina Island, November 1991—December 1992. A re-

port to the Damage Assessment Office, U. S. Fish and Wildlife Service, Sacramento, California. PPX007

4146. 24 pp.

. 1994c. Food habits of bald eagles on Santa Catalina Island, January — July 1993. A report to the Damage
Assessment Office, U. S. Fish and Wildlife Service, Sacramento, California. PPX007 4171. 17 pp.

GARCELON, D. K., R. W. RISEBROUGH, W. M. JARMAN, A. B. CHARTRAND, AND E. E. LITTRELL. 1989. Accumu-
lation of DDE by bald eagles reintroduced to Santa Catalina Island in Southern California. Pp. 491-494 in
Proceedings of the Third World Conference on Birds of Prey and Owls, Israel.

GILMARTIN, W.G., R. L. DELONG, A. W. SMITH , J. C. SWEENEY, B. W. DE LAPPE, R. W. RISEBROUGH, L. A.
GRINER, M. D. DAILEY, AND D. B. PEAKALL. 1976. Premature parturition in the California sea lion. Journal
of Wildlife Diseases 12:104—114.

HANAN, D. J. 1996. Dyamics of abundance and distribution for Pacific harbor seals, Phoca vitulina richardsi, on
the coast of California. Ph.D. Dissertation, University of California, Los Angeles. 158 pp.

HANAN, D., J. SCHOLL, AND S. DIAMOND. 1986a. Harbor seal, Phoca vitulina richardsi, census in California,
June 25-30, 1984. Pp. 2-7 in D. A. Hanan, California Department of Fish and Game Coastal Marine Mam-
mal Study. Annual report for the period July 1, 1983—June 30, 1984. NOAA/NMFC SWEC Admin. Rep.
LJ-86-16C; 55 pp.

LE BOEUF: PINNIPEDS ON SANTA CATALINA ISLAND 21

. 1986b. Harbor seal, Phoca vitulina richardsi, census in California, May 28-31 and June 25—30, 1985.
Pp. 2-8 in California Department of Fish and Game Coastal Marine Mammal Study. Annual report for the
period July 1, 1984—June 30, 1985. D. A. Hanan. NOAA/NMFC SWFC Admin. Rep. LJ-86-25C. 46 pp.

. 1987. Harbor seal, Phoca vitulina richardsi, census in California, June 2—5, 30 and July 1, 1986. Pp. 2-8

in NOAA/NMFC SWEC Admin. Rep. SWR-87-3. 41 pp.

. 1988. Harbor seal, Phoca vitulina richardsi, census in California, May—June, 1987. NOAA/NMFC
SWFC Admin. Rep. SWR-88-2. 49 pp.

Kirr, L. F. 1980. Historical changes in resident populations of California islands raptors. Pp. 65 1—653 in The Cal-
ifornia Islands: Proceedings of a Multidisciplinary symposium. Santa Barbara Museum of Natural History,
D. M. Power, ed. Santa Barbara, California. 787 pp.

LE Boeur, B. J. AND M. L. BONNELL. 1971. DDT in California sea lions. Nature 234:108—110.

LIEBERG-CLARK, P., C. E. BACON, S. A. BURNS, W.M. JARMAN, AND B. J. LE BOEuF. 1995. DDT in California
sea-lions: A follow-up study after twenty years. Marine Pollution Bulletin 30:744—745.

Lowry, M.S. ANDR. L. FOLK. 1987. Feeding habits of California sea lions from stranded carcasses collected at
San Diego County and Santa Catalina Island, California. National Marine Fisheries Service, Southwest
Fisheries Center, P.O. Box 271, La Jolla, California 92038, Administative Report LJ-87-15. 22 pp.

Lowry, M. S., P. BOVENG, R. L. DELONG, C.W. OLIVER, B. S. STEWART, H. DEANDA, AND J. BARLOW. 1992.
Status of the California sea lion (Zalophus californianus californianus) population in 1992. Southwest Fish-
eries Science Center, Administrative Report LJ-92-32. 24 pp.

MATE, B. R. 1977. Aerial censusing of pinnipeds in the eastern pacific for assessment of population numbers, mi-
gratory distributions, rookery stability, breeding effort, and recruitment. Final Rept. MMC-75/01 U. S. Mar.
Mamm. Comm., Washington, D. C. NTIOS PB-265 859. 67 pp.

MACGREGOR, J. S. 1974. Changes in the amount and proportions of DDT and DDD, in the marine environment
off southern California, 1949-72. U.S. National Marine Fisheries Service Fisheries Bulletin
72(2):275-293.

ODELL, D. K. 1971. Censuses of pinnipeds breeding on the California Channel Islands. Journal of Mammalogy
52(1):187-190.

PETERSON, R. S. AND G. A. BARTHOLOMEW. 1967. The Natural History and Behavior of the California Sea Lion.
Special Publication No. 1, The American Society of Mammalogists.

RipLey, W. E.,K. W. Cox AND J. L. BAXTER. 1962. California sea lion census for 1958, 1960 and 1961. Califor-
nia Fish and Game 48:228—231.

ROWLEY, J. 1929. Life history of the sea-lions on the California coast. Journal of Mammalogy 10:1—36.

SEAGARS, D. J., D. P. DEMASTER, AND R. L. DELONG. 1985. A survey of historic rookery sites for California
and northern sea lion in the Southern California Bight. Southwest Fisheries Center Administrative Report
No. LJ-85-13. 61 pp.

SHAW, S.B. 1971. Chlorinated hydrocarbon pesticides in California sea otters and harbor seals. California Fish
and Game 57:290—294.

STEWART, B. S. 1982. Peak 1982 aerial censuses of harbor seal populations on the Southern California Channel
Islands. Hubbs Sea World Research Institute Tech. Rep. 82-143. 7 pp.

WEISE, M. J. 1999. Abundance, food habits, and annual fish consumption of California sea lion (Zalophus
californianus) and its impact on salmonid fisheries in Monterey Bay, California. M. S. thesis. San Jose State
University, San Jose, California. 102 pp.

PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENC

LIDIALY

JUN 20 2002 _ Jpne 14, 2002

Volume 53, No. 3, pp. 23-36, 5 figs., 1 pl., 2 tables.

Wobea Hole, MA 02543
A Revision of Pachydactylus scutatus (Reptilia: Squamata:
Gekkonidae) with the Description of a New Species from
Northern Namibia

by
Aaron M. Bauer!,Trip Lamb’, and William R. Branch’

| Department of Biology, Villanova University, S00 Lancaster Avenue,
Villanova, Pennsylvania 19085; email: aaron.bauer@villanova.edu
“Department of Biology, East Carolina University,
Greenville, North Carolina, 27858; email: lamba@mail.ecu.edu
3Port Elizabeth Museum, P.O. Box 13147, Humewood 6013, Republic of South Africa
email: pemwrb@zoo.upe.ac.za

A review of taxa previously synonymized with Pachydactylus scutatus (Gekkonidae) re-
veals that both P. angolensis and P. robertsi should be regarded as specifically distinct. Al-
though the former species appears to be closely related to P. scutatus, the latter is probably
more closely allied to the P. weberi complex. A new member of the P. scutatus group is de-
scribed from the Kaokoveld of northern Namibia. Both diagnostic morphological features
and DNA sequence divergence observed for the cytochrome b gene support the distinctive-
ness of the new taxon.

The genus Pachydactylus is the most speciose group of geckos in southern Africa. At least 34
species are currently recognized (Kluge 2001), 20 of which occur in Namibia (Branch 1998; Griffin
2000). Although the genus is well represented throughout Namibia, the country’s rocky, arid north-
west is especially species rich (Bauer et al. 1993; Griffin 2000). The last complete revisions of
Pachydactylus (FitzSimons 1943; Loveridge 1947) were conducted before many taxa from this re-
gion were first collected. Increased field work in northwestern Namibia from the 1960s onwards re-
sulted in the discovery of species previously known only from Angola (e.g., Pachydactylus
caraculicus; Haacke 1970) and in the description of several new taxa including Pachydactylus
oreophilus (McLachlan and Spence 1967), P. gaiasensis, and P. sansteyni (Steyn and Mitchell 1967).
Subsequent revisions and systematic analyses of subgroups of Pachydactylus (e.g., McLachlan and
Spence 1966; Broadley 1977; Bauer and Branch 1995; Branch et al. 1996; Lamb and Bauer 2000, in
press) have not addressed the phylogeny or taxonomy of the northern Namibian species.

One of the most distinctive geckos of northern Namibia, Pachydactylus scutatus, was described
from Kowaris [= Kowares] near Etosha by Hewitt (1927), who considered it closely related to P.
montanus (a synonym of P. serval). FitzSimons (1938) reported additional specimens from
Kamanjab, Paderborn, and Huab, all to the south of the type locality. In the same paper, FitzSimons
also described Pachydactylus robertsi from a unique holotype from Farm Kraikluft in the Great Karas
Mountains. FitzSimons noted that P. robertsi superficially resembled P. scutatus, but could be distin-
guished by nostril position, dorsal and caudal scalation, and head color.

Subsequently, Loveridge (1944) described P. scutatus angolensis from Hanha and Lobito Bay in
Benguella Province, Angola. He also re-evaluated the status of P. robertsi, reducing it to a subspecies

24 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES
Volume 53, No. 3

of P. scutatus, even though he had never examined specimens of either P. scutatus or P. robertsi. He
regarded all three forms as “‘a further development of the scaly-backed P. p. punctatus” and based his
subspecific asignments on the continuity of variation in dorsal keeling among them. Although
Loveridge’s taxonomy has received broad support (e.g., Loveridge 1947; Mertens 1955; Wermuth
1965: Kluge 1993, 2001; Résler 2000), some recent herpetological works dealing with Namibian
Pachydactylus (e.g., Branch 1998; Bauer 1999[2000]) have not recognized subspecies within P.
scutatus.

Mertens (1955), Van den Elzen (1983), and Bauer and colleagues (1993) reported numerous ad-
ditional localities for Pachydactylus scutatus, but P. robertsi remains known only from the unique
holotype, and only four additional specimens of P. s. angolensis, all from near Namibe, Angola
[Quarter Degree Square (QDS) 1512Aa: 15° latitude, 12° longitude, upper case letters identify quad-
rants, lower case letters quadrants within quadrant], have been reported in the literature (Laurent
1964). Laurent (1964) provided some scale counts for these specimens but, unfortunately, none that
are informative within the P. scutatus group. These specimens, catalogued in the Dundo collection in
Angola, are not currently accessible, and ongoing civil strife in Angola precludes the collection of
fresh material. Due in part to the paucity of material, a critical evaluation of the affinities and validity
of the non-nominate members of the P. scutatus group has yet to be undertaken. Indeed, the only pub-
lished statement concerning phylogenetic relationships within this group is that of FitzSimons (1959),
who believed that Pachydactylus caraculicus was perhaps both geographically and phylogenetically
intermediate between P. s. angolensis and P. s. scutatus. This interpretation was challenged by
Laurent (1964), who noted that P. s. angolensis occurs in sympatry with P. caraculicus in southern
Angola and, therefore, must be specifically distinct.

As part of a major revision of Pachydactylus and related geckos, we re-examined the types of P.
robertsiand P. s. angolensis in light of current knowledge of the distribution and morphological varia-
tion of P. s. scutatus. We also collected additional specimens of of the P. scutatus group from the
Kaokoveld and adjacent parts of the Kunene Region of Northern Namibia for DNA sequence analysis
to determine if morphological variation within the group is reflected by genetic divergence. We here
present a revision of the P. scutatus group, which entails the revalidation of P. robertsi and P.
angolensis as full species, as well as the description of a new taxon from extreme northwestern
Namibia.

MATERIAL AND METHODS

The following measurements were taken with Brown and Sharpe Digit-cal Plus digital calipers
(to the nearest 0.01 mm): snout-vent length (SVL; from tip of snout to vent), tibia length (TBL; from
base of heel to knee); tail length (TL; from vent to tip of unregenerated tail), tail width (TW; measured
at base of tail); head length (HL; distance between posterior edge of last supralabial and snout-tip),
head width (HW; measured at angle of jaws), head depth (HD; maximum height of head, from occiput
to throat), ear length (EL; longest dimension of ear); forearm length (FA; from base of palm to el-
bow); eye diameter (ED; greatest diameter of orbit), eye to nostril distance (E-N; distance between
anteriormost point of eye and nostril), eye to snout distance (E-S; distance between anteriormost point
of eye and tip of snout), eye to ear distance (E-E; distance from anterior edge of ear opening to poste-
rior corner of eye), and interorbital distance (IO; shortest distance between medial rims of left and
right orbits).

Scale counts and external observations of morphology were made using a Nikon SMZ-10 dis-
secting microscope. Radiographic observations were made using a Faxitron closed cabinet X-ray sys-
tem. Comparisons were made with museum material, as well as original published descriptions and
descriptions provided in broader faunal and taxonomic treatments. Standard codes for museum col-
lections follow Leviton et al. (1985).

BAUER, LAMB, AND BRANCH: PACHYDACTYLUS SCUTATUS 25

DNA sequence analysis was also conducted to determine whether morphological features distin-
guishing the new form from P. scutatus and other congeners were complemented by substantive ge-
netic divergence. Tissue samples from nine specimens (six members of the P. scutatus group and
three representing two other congeners) were processed at field collection sites and preserved in a sat-
urated salt-DMSO buffer (Amos and Hoelzel 1991). Genomic DNA was extracted from liver using
the Qiagen QIAamp DNA Mini kit. Portions of the mitochondrial cytochrome b (cyt b) gene were
used to assess sequence variation among specimens. The primers L14724 and H15149 (Meyer et al.
1990) were used to amplify a 400 bp segment of the cyt b gene; 50 ul reactions were amplified for 32
cycles at 92°C for 45 sec, 55°C for 35 sec, and 72°C for 1 min. Amplification products were purified
over Centri-sep columns and served as templates in cycle-sequencing reactions employing
dye-labeled terminators (PRISM kit, Applied Biosystems, Inc.). PRISM reaction products were ana-
lyzed on an Applied Biosystems 373A automated DNA sequencer. Forward and reverse sequences
were generated for each sample and their complementarity confirmed using the Sequence Navigator
software (Applied Biosystems, Inc.). Sequences were aligned using the CLUSTAL X program, ap-
plying default settings (Thompson et al. 1997). Pairwise sequence divergence estimates were derived
using the Hasegawa-Kishino-Y ano model (HK Y85; Hasegawa et al. 1985), which assumes different
evolutionary rates for transitions and transversions and unequal base frequencies.

SYSTEMATICS

Examination of the types of P. robertsi and P. scutatus angolensis in light of current understand-
ing of the geographic range and morphological variation of typical P. scutatus suggests that both taxa
should be considered specifically distinct. FitzSimons (1938) and subsequent workers recognized the
affinity of P. robertsi to P. scutatus on the basis of enlarged dorsal scales and a distinct, white nuchal
collar.

The unique type of P. robertsi (Fig. 1) possesses moderately well-developed keels on enlarged,
weakly imbricate dorsal scales. Although the dorsal scales superficially resemble those of P. scutatus,
most other characteristics indicate an affinity with the Pachydactylus weberi complex (including P.
weberi sensu stricto and several taxa currently accorded subspecific status or regarded as synonyms of
P. weberi). In particular, the exclusion of the rostral, but not the first supralabial, from the nostril typi-
fies P. weberi, and the degree of dorsal scale keeling falls within the range of Namibian specimens of
this species. In addition to scalation, similarities in the nuchal collar of P. robertsi have been consid-
ered evidence for its affinities with P. scutatus. However, adults of many members of the P. weberi
group commonly retain elements of the juvenile pattern, which most frequently consists of three
bands: one on the neck, one near mid-body and one in the sacral region. In P. robertsi and certain P. cf.
weberi, especially those from central Namibia (e.g., Windhoek: TM 32834, 328383; Klein
Windhoek: TM 28781; Horebis Nord, Karibib District: TM 33650), only the anterior band is retained.
The white collar in P. scutatus is usually one scale width thick (Fig. 5), whereas in P. robertsi it spans
2-3 scales and is situated somewhat more posteriorly on the neck (Fig. 1).

The assignment of P. robertsi to the P. weberi complex also makes sense geographically. At the
time of the description of P. robertsi, P. scutatus was known from only four localities. Based on mod-
ern collections, the range of P. scutatus is strictly limited to the region from the Brandberg to the north
and west, more than 700 km from the P. robertsi type locality. Pachydactylus robertsi is quite distinc-
tive from typical P. weberi from Little Namaqualand, but its relationships within the broader context
of the P. weberi complex as a whole remain unresolved. At present we regard P. robertsi as a distinc-
tive species allied to P. weberi and tentatively restricted to the Great Karas Mountains. This taxon will
be considered in more detail in a revision of the P. weberi complex (Bauer et al., in prep.).

Pachydactylus s. angolensis (Fig. 2) is clearly more similar to P. s. scutatus than is P. robertsi.
Loveridge (1944, 1947) noted the exclusion of both the first supralabial and rostral scales from the

26 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES
Volume 53, No. 3

nostril in this form. This condition, the chief di-
agnostic feature employed in the taxon descrip-
tion, presumably also characterizes the four
additional specimens reported on by Laurent
(1964), who noted nothing to the contrary.
Loveridge’s types were collected at Hanha
(probably Hanha do Norte in quarter degree
square 1213Bc another locality, Hanha do
Cubal, 1314 Aa is also in the same province) and
Lobito Bay (1213Bc), both in Benguela Prov-
ince. Laurent’s (1964) material was from the
“environs de Mocamedes” (now Namibe,
Namibe Province), approximately 335 km south
of the type locality. The rostral/supralabial scale
condition has not been recorded in any of the
typical P. scutatus examined by us or reported in
the literature. As this character appears to be
constant, we regard it as diagnostic evidence for
the recognition of P. angolensis as distinct from
P. scutatus. These taxa also differ in color pat-
tern and dorsal and caudal scalation (outlined in
the morphological comparisons section of the
Discussion below).

The occurrence of P. angolensis in coastal

and near coastal localities also suggests ecologi-
cal differences from P. scutatus. Laurent’s FIGURE |. Holotype of Pachydactylus robertsi (TM 17854).
(1964) specimens were collected from crevices Note the position and width of the nuchal collar. Scale
in friable rock along the roadside from Sa de >> !0™™-
Bandeira to Namibe and under rocks on the littoral plateau of “das Conchas” beach, whereas typical P.
scutatus are usually associated with rock flakes on exfoliating boulders (FitzSimons 1938; Bauer et al.
1993). On the basis of its distinctive scalation, coloration, and ecology, we regard Pachydactylus
angolensis as a valid species, probably closely allied to P. scutatus. At present all known specimens
are from lowland, near coastal localities in Namibe and Benguela provinces, southwestern Angola.
The southernmost locality, Namibe (1512Aa), is approximately 250-280 km northwest of the north-
ernmost localities of P. scutatus (Otjivize, 1713Cb and Epupa, 1613Cd), but the intervening area has
been poorly collected and the true extent of the distribution of P. angolensis remains uncertain.

Even upon the removal of P. robertsi and P. angolensis from synonymy with P. scutatus, signifi-
cant morphological variation among populations of P. scutatus remains. Specimens we collected near
Sesfontein, as well as museum material from Kaoko Otavi and Orupembe, are especially distinctive.
Based upon detailed comparisons with P. scutatus and P. angolensis, we here recognize and describe
a third member of the P. scutatus group.

REPTILIA: SQUAMATA: GEKKONIDAE

Pachydactylus parascutatus sp. nov.
Fig. 331Ply i

HOLOTYPE. — National Museum of Namibia (SMW) R 9454 (previously California Academy
of Sciences [CAS] 214755; Field Number AMB 6518) (adult female), from the vicinity of Para Camp,

BAUER, LAMB, AND BRANCH: PACHYDACTYLUS SCUTATUS Pal

2km north of Sesfontein, Opuwo Dis-
trict, Kunene Region, Namibia
mi8707'28"S, 13°35'29"E), collected by
R. D. Babb, A. M. Bauer, T. Lamb, W. R.
Branch, P. Moler, V. Schneider, S.
Lochetto, and A. Meeker, 3 June 2000.

PARATYPES. — CAS 214735 (Field
number AMB 6498; adult male), same lo-
cality and collectors as holotype, col-
lected 2 June 2000; CAS 214750 (Field
number AMB 6512; adult female),
Purros Rd., ca. 8 km west of Sesfontein,
Opuwo District, Kunene Region,
Namibia (19°10'05"S, 13°34'03"E), col-
lected by R. D. Babb, A. M. Bauer, T.
Lamb, W.R. Branch, P. Moler, V.
Schneider, S. Lochetto, and A. Meeker, 3
June 2000: SMW R 182 (adult male),
Kaoko Otavi, Opuwo District, Kunene
Region, Namibia, collected by W. Steyn,
8 October 1965; SMW R 184 (adult fe-
male), 20 km west of Orupembe, Opuwo
District, Kunene Region, Namibia, col-
lected by W. Steyn, 6 October 1965.

ADDITIONAL MATERIAL
EXAMINED. — CAS 214749 (Field num- Figure 2. Holotype of Pachydactylus angolensis (AMNH R 47874).
ber AMB 6511; juvenile), 27 km, Purros Scale bar = 10 mm.

Rd., ca. 27 km W of Sesfontein, Opuwo
District (19°03'16"S, 13°30'08”E), collected by R. D. Babb, A. M. Bauer, T. Lamb, W. R. Branch, P.
Moler, V. Schneider, S. Lochetto, and A. Meeker, 3 June 2000.

ETYMOLOGY. — The specific epithet reflects the similarity of the new species to P. scutatus and
also commemorates the type locality near “Para Camp,” a rustic camp site operated by villagers near
the town of Sesfontein.

DIAGNOSIS. — Snout-vent length to 38.40 mm (largest female paratype); body sub-cylindrical,
not depressed; most dorsal scales large, rounded, strongly keeled, juxtaposed or weakly imbricate, in
approximately 14 rows; ventral scales subimbricate, scales on dorsum of forelimb and granules be-
tween larger dorsal scales unkeeled; nostril surrounded by two postnasals, supranasal, rostral and first
supralabial; enlarged scansorial lamellae under digit IV of pes 4-5; adult dorsal pattern with numer-
ous dark brown markings, without distinct white nuchal collar; original tail more-or-less banded;
venter buff with light brown speckling.

DESCRIPTION (based on holotype). — Adult female (Fig. 3, Pl. 1). Snout-vent length 35.25 mm.
Body sub-cylindrical, not markedly depressed, elongate (A-G/SVL ratio 0.44). Head elongate, large
(HL/SVL ratio 0.29), moderately wide (HW/SVL ratio 0.19), somewhat depressed (HD/HL ratio
0.36), distinct from neck. Lores and interorbital region inflated. Snout short (E-S/HW ratio 0.57), lon-
ger than eye diameter (ED/E-S ratio 0.63); scales on snout and forehead flattened, round to oval;
scales on snout much larger than those of parietal table. Eye relatively small (ED/HL ratio 0.23); or-
bits without extra-brillar fringes; posterior supracilliary scales bearing small spines; pupil verti-
cal,with crenelated margins. Ear opening oval, small (EL/HL ratio 0.07), greatest diameter horizontal;
eye to ear distance greater than diameter of eyes (E-E/ED ratio 1.09). An irregular series of flattened to

28 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES
Volume 53, No. 3

weakly conical tubercles between posterior border of orbit and occiput. Rostral approximately half as
deep as wide; no rostral groove; contacted by two enlarged supranasals and first supralabials; nostrils
rounded, each surrounded by two postnasals, supranasal, rostral, and first supralabial; supranasals in
broad contact anteriorly, separated posteriorly by a single small granule; dorsal postnasals separated
by two granules anteriorly and three posteriorly; one row of scales separate orbit from supralabials.
Mental rectangular, approximately three times deeper than wide; no enlarged postmentals or chin
shields.

Scales grade from granular on parietal region to a mixture of granules and enlarged conical tuber-
cles on occiput, becoming keeled on nape; dorsal body scales large, rounded, flattened, with a pro-
nounced median keel, forming approximately 14 rows; a single granular scale present in the space
formed at the junction of four adjacent keeled tubercles (Fig. 4A); scales decreasing sharply in size on
flanks, becoming smooth, subimbricate; ventral scales flattened, subimbricate, becoming somewhat
larger posteriorly; scales on dorsum at mid-body much larger than those on ventrum at same level;
chin granules approximately one third to one half size of ventral scales, increasing gradually in size on
throat. No preanal or femoral pores; no preanal groove. Scales on palm and sole smooth, rounded;
scales on ventral aspect of hind limbs and dorsal aspect of forelimbs imbricate, not keeled; those on
dorsal aspect of distal thigh and crus greatly enlarged, keeled to mucronate.

Forelimbs moderately short, stout; forearm short (FA/SVL ratio 0.11); hindlimbs relatively
short, tibia short (TBL/SVL ratio 0.15); digits moderately short, minute claws present on digits II and
III of the pes only; subdigital scansors, except for distalmost, entire, present only on distal portion of
toes, 1.5—2.0 times wider than more basal (non-scansorial) subdigital scales; interdigital webbing ab-
sent. Relative length of digits (manus; measurements in mm in parentheses): IV (1.71) > UI.(1.70) > II
(1.68) > V (1.60) >1 (1.49); (pes): IV (3.02) > III (2.61) > V (2.50) > 1 (1.95) >1
(1.08). Subdigital scansors (manus): I (3), II (4), HI (3), IV (3), V (3); (pes) I
G4) G), TG) sIVeG); VG):

Tail cylindrical; partially regenerated tail longer than snout-vent length
(TL/SVL ratio 1.13); tail relatively thick, tapering, with distinct whorls of
scales; each transverse row of enlarged, keeled tubercles separated by two
rows of smaller scales; smaller scale rows continuous around tail; each row of
enlarged tubercles replaced ventrally by two rows of smaller scales; each row
of keeled dorsal caudal tubercles more-or-less continuous, interrupted occa-
sionally by isolated small granules; corresponding to two rows of ventral
scales; subcaudal scales pointed posteriorly, imbricating; two somewhat en-
larged, smooth, conical postcloacal spurs on each side of tailbase.

Scutellation. Supralabials to angle of jaws 8; infralabials 7; interorbital
scale rows (at midpoint of orbit) 19 (8 across narrowest point of frontal bone);
midbody scale rows at belly to lowest row of keeled dorsal tubercles 37.

Osteology. Premaxillary teeth 9; phalangeal formulae 3-3-4-5-3 for
manus and 3-3-4-5-4 for pes; presacral vertebrae 26, including three anterior
cervical (without ribs) and one lumbar; sacral vertebrae 2; 5 pygal and 8.5 post
pygal caudal vertebrae to point of regeneration; no cloacal bones present.
Endolymphatic sacs enlarged, filled with calcium.

Coloration (in life; Pl. 1). Ground color of dorsum beige to yellowish
brown with relatively regularly arranged mid-brown, roughly diamond-shaped
markings; flanks with a pale rosy flush from underlying tissue; original tail
with a series of darker markings at each pair of smaller dorsal scale rows, form-

FiGuRE 3. Holotype of P. parascutatus, sp. nov. (SMW R 9454) from Para Camp, 2 km north of Sesfontein, Opuwo District,
Namibia. Scale bar = 10 mm.

BAUER, LAMB, AND BRANCH: PACHYDACTYLUS SCUTATUS 29

PLATE |. Holotype of Pachydactylus parascutatus sp. nov. in life.

ing incomplete bands on some tail segments; dorsal surfaces of limbs mottled with scattered brown
markings similar to those of dorsum. A thick, brown line from nostril, across canthus rostralis, to orbit.
continuing posteriorly to approximately level of occiput, breaking up into markings that merge with
dorsal pattern: bordered above by a thick whitish line, most distinct between nostril and anterodorsal
corner of orbit. A series of three weakly defined, mid-brown. concentric semicircles extending poste-
riorly from frontoparietal suture. posterior border of orbit and occiput, respectively. Labial scales
whitish; anterior supralabials and all infralabials bearing mid-brown markings. Venter buff with light
specklings of brown pigment on all but midventral scales of chest and abdomen, densest on underside
of tail and thighs.

VARIATION. — The adult paratypes (Table 1) are similar to the holotype in most respects, but dif-
fer in the following ways: CAS 214750 — dorsal pattern bolder, forming more regular longitudinal
markings, especially on nape and shoulders: CAS 214735 — band from eye continues on left side be-
yond shoulder. dorsal markings fused. but still irregular, hemipeneial bulge present, two large white
pointed cloacal spurs on each side of vent, SMW R 182—more strongly patterned than holotype with
darker brown markings, three clocal spurs per side, precloacal scales larger than abdominal scales:
SMW R 184—lacking distinct white canthal stripe. head markings more diffuse. virtually no ventral
speckling (perhaps an artifact of blanching in preservative), four cloacal spurs per side. In the single
juvenile specimen examined (CAS 214749), the white canthal lines continue behind the orbits, fusing
on the nape to forma distinctive V-shaped collar as in P. scutatus. Males possess a single pair of cres-
centic cloacal bones. similar to those of P. scutatus and P. angolensis, and have prominent hemipenial
bulges and somewhat enlarged cloacal spurs. We could find no evidence of even minute claws in ei-
ther of the male specimens examined.

DISTRIBUTION. — The specimens examined are derived from a small area in the Kaokoveld
north of the Hoanib River. All localities are relatively arid, but the two eastern areas, near Sesfontein
and Kaoko Otavi are characterized by open mopane woodland or savanna, whereas the locality to the
west of Orupembe is dominated by grasses and other herbaceous plants and is considered part of the
northern Namib proper (Giess 1971). The species thus occurs in two adjacent floral domains or
biomes (Jiirgens 1991; Irish 1994). PEM R1964—5 from “Sesfontein” appear to be typical P. scutatus.
Although this locality. if accurate. would imply sympatry with P. parascutatus, it is probable that it is
imprecise and that the specimens may have been derived from some distance to the south of
Sesfontein itself, within the accepted range of typical P. scutatus. Typical P. scutatus are also known
from at least two localities (Otjivize and Epupa) to the north of P. parascutatus. In order to determine

30 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES
Volume 53, No. 3

FiGure 4. Mid-dorsal back scalation of A) Pachydactylus parascutatus and B) P. angolensis. Note the small, mostly granular
scales at the junction of each quartet of keeled imbricate scales in A in comparison to their larger, often keeled counterparts in B.

whether P. parascutatus actually occurs in strict sympatry with either P. scutatus or P. angolensis, all
older museum material should be re-examined and additional collecting should be conducted in the
area from Sesfontein north to the Kunene River and into southern Angola.

NATURAL HISTORY. — The holotype was collected at night (approximately 20 hr) active on the
surface of limestone rocks ona hillside. CAS 214735 was collected on the same hillside. CAS 214750
was collected by day amongst rubble beneath a large, flat slab of sandstone near the top ofa steep slope
beneath an overhanging cliff. CAS 214749 was collected by day from a vertical crevice between two
large flat pieces of sandstone. In contrast Pachydactylus scutatus typically shelters under rock flakes
on exfoliating boulders (FitzSimons 1938; Bauer et al. 1993), or occasionally in deeper rock cracks.
The fact that P. scutatus have been taken in pitfall traps (SMW R 7972, 8002, 8032), however, sug-
gests that this species may also be active terrestrially and not as strictly rupicolous as generally be-
lieved. Based on the capture conditions of the types of P. parascutatus, the microhabitat preference of
this species appears intermediate between P. scutatus and P. angolensis, which has been taken from
crevices in friable rocks and from under stones on the ground (Laurent 1964).

BAUER, LAMB, AND BRANCH: PACHYDACTYLUS SCUTATUS 31

TABLE |. Mensural data for the type series of Pachydactylus parascutatus, sp. nov. Abbreviations as in
Materials and Methods, all measurements in mm.

Holotype Paratypes
SMW R 9454 CAS 214735 CAS 214750 SMW R 182 SMW R 184

Sex Female Male Female Male Female
SVL goes 33.58 37210 34.56 38.40
FA 4.02 4.96 13 4.70 4.48
TBE 5.43 4.96 5.40 5.61 5.84
TL (total) 26.30 9.67 33.89 25.16 3.63
TL (regen.) 12.86 tail broken 20.02 [ies Sel tail broken
TW 3.23 DOD Bo 359 3.86
A-G 15.41 14.07 17.04 13.44 16.95
HL 10.31 9.84 11.78 9.99 9.84
HW 6.43 5.83 6.05 6.23 6.59
HD 3.76 3.04 3.61 4.02 3.37,
ED Vip! 2.06 2.40 2.42 2.38
E-E psy) pe he 2.90 2.89
E-S 3.68 3.62 3.69 So 3.03
E-N 3.2 2.65 2.56 2.41 E32
IO 2.88 z.19 2:79 3.30 2.70
EL 0.72 0.67 0.84 0.62 0.64

Like that of P. scutatus, the skin of P. parascutatus 1s fragile and easily damaged by handling
(Bauer and Russell 1992; Bauer et al. 1993). The skin of the flank and limbs of several paratypes (CAS
214735, 214750) were damaged in collection. Fragile skin in these forms is a manifestation of re-
gional integumentary loss, an antipredator escape strategy that has been proposed as useful against
certain types of lizard predators, such as snakes and other lizards (Bauer et al. 1989, 1993; Bauer and
Russell 1992).

Other reptiles collected at Para Camp, Sesfontein, were Pachydactylus oreophilus, P. turneri,
Rhoptropus barnardi, Leptotyphlops occidentalis, Prosymna visseri, and Pythonodipsas carinata.
Psammophis trigrammus was collected at one of the paratype localities.

DISCUSSION

MORPHOLOGICAL COMPARISONS. — The relatively small body size (to 38.40 mm) of P.
parascutatus easily distinguishes it from P. tuberculosus, P. tetensis, and all members of the P.
bibronii, and P. namaquensis groups. These are basal members of the genus (Lamb and Bauer 2002),
all of which exceed 100 mm in adult SVL and possess 10 or more lamellae beneath the fourth toes of
the pes (vs. 4-5 in P. parascutatus). Among “small-bodied” Pachydactylus, the combination of en-
larged, contiguous, keeled dorsal scales and the entry of both the first supralabial and the rostral into
the nostril occurs only in the P. scutatus group. Some members of the P. weberi complex (only one
species currently recognized [Branch 1998], but currently under revision) have variably developed
keels and, although the first supralabial typically enters the nostril, the rostral is usually excluded.
Only in the case of P. robertsi (see above) do the dorsal scales approach the width of those of P.
parascutatus, but the rostral is excluded from the nostril in P. robertsi.

Within the Pachydactylus scutatus group, P. parascutatus differs from P. angolensis in the con-
dition of the nasal scales (both rostral and supralabial excluded from nostril in P. angolensis). In addi-
tion, in P. angolensis the small scales lying at the junction of each quartet of larger dorsal scales are
frequently keeled (Fig. 4B). These small scales are flat or granular in the new species (Fig. 4A). The
large dorsal scales are relatively strongly imbricate in P. angolensis, but weakly so or juxtaposed in P.

o>)
bo

PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES
Volume 53, No. 3

parascutatus. In addition, the former species has keeled scales on the dorsal surface of the forelimb
(vs. smooth) and small, sometimes keeled scales separating the enlarged caudal tubercles (vs. no or
few small intervening granules). Pachydactvlus angolensis also has keeling on the rows of smaller
caudal scales (between tubercular whorls), whereas such keeling is weak or altogether absent in P.
parascutatus. The dorsal pattern may also be diagnostic. Loveridge (1944) described the types of P.
angolensis as grayish to reddish brown and noted the presence of white spots on the dorsum. At the
time of Loveridge’s description, the types had already been in preservative for 19 years, and the life
colors were clearly highly modified. The pattern of the holotype today 1s at best weakly discernable.
However, although similar head patterning is evident, there is no evidence of the dark dorsal markings
typical of P. parascutatus. A paratype of P. angolensis (MCZ 49856) exhibits no evidence of a neck
band and possesses remnants of broken transverse bands formed by pale scales on the dorsum.
Pachydactylus parascutatus shares with P. scutatus the same snout scalation pattern and is very
similar in most aspects of body scalation. It differs from this form chiefly in color pattern. Adult P.
parascutatus lack the well-defined nuchal collar that typifies P. scutatus (Fig. 5). However, this collar
is evident in the single juvenile specimen (CAS 214749) of P. parascutatus that we examined. Even in
this specimen, however, the collar lacks the dark edging characteristic of P. scutatus. In P.
parascutatus the light head markings are less well defined than in P. scutatus, where they are often 1m-
maculate and bold. In addition, most P. scutatus exhibit a uniform or very weakly patterned dorsum
(Fig. SA; although dark scattered markings do occur in some specimens, e.g., SMW R 5183 and CAS

FIGURE 5. Pachydactylus scutatus from A) Farm Franken, Outjo District, Kunene Region, Namibia (CAS 176081) near the
type locality of Kowares and B) 19°37'27"S, 14°43'19"E, Khorixas District, Kunene Region, Namibia. The unpatterned dor-
sum of A is the most common pattern. B is a particularly heavily patterned individual. Compare the position and width of the
nuchal collars of these specimens with the type of P. angolensis (Fig. 1). Scale bars = 10 mm.

BAUER, LAMB, AND BRANCH: PACHYDACTYLUS SCUTATUS 33

a 193673; Fig. 5B; see also Branch 1998, plate 85)
a ¢ with a unicolor or diffusely-patterned tail, whereas
2 ) = P. parascutatus are relatively heavily patterned,
on with dark marks over most of the dorsum and a dis-
AS! 3
g ns tinctly banded tail. Pachydactylus scutatus also ap-
Coo tT :
nt ~ +O pear to grow to much larger sizes than the new
> ees ee e
= = a species. Branch (1998) recorded a maximum size
ie of 42 mm SVL for P. scutatus, but the largest spec-
9 aon imens we examined attained sizes of approxi-
eh 0 | ess mately 47 mm SVL, in comparison to a maximum
S Peso of less than 39 mm in P. parascutatus. Radio-
2 | | graphs of specimens of P. scutatus of 33-38 mm
g | | sae SVL reveal that they are juveniles or subadults,
3 wm S288 with poor ossification of the limb joints, whereas
S xe yes P. parascuatus in this size range are adults. The
z largest of the types of P. angolensis measures
= | OO at 42 mm SVL. Pachydactylus scutatus also has a
2 mmm 03 mM ‘ 2 :
= ;
= i ee SPs somewhat more flattened or depressed habitus
~ 'SSHAQQ : :
S Ssocco than the new species.
= MOLECULAR COMPARISONS. — Pairwise
~— .
8 2
a Seo CON sie comparisons between Pachydactylus parascutatus
3 vi SSSESESE and P. scutatus yielded genetic distance values for
= hSeee=agag
8 Seeceoceose cytb in the range of 17.6-20.3%, whereas
N intraspecific comparisons reveal differences of
= p p
S S S oo 23 eS os 0.0-0.5% among P. parascutatus (maximum geo-
fon] No) oO : ‘ ;
5a AIRS RS graphic distance between samples approximately
Z a ooooooo 25 km) and 5% between P. scutatus from across a
a | 35 km distance (Table 2). These interspecific com-
é 3. S S S % 2 “Ss parison values are only slightly less than those be-
2 S | l 5 Aaag a Fs = ae RP: Si esate a P. sansteyni (21.7-
= 5 Seoooasce 5%), another Kaokoveld regional endemic spe-
oO vO : .
ae <t cies, and P. weberi (20.3—20.6%), another strongly
Ong nod a 4
eS ae t oot & ae Ms keeled member of the genus. Like most interspe-
eee) eo Se Se oy cific comparisons in Pachydactylus, these values
Soe! El anwtanaanae”a : ey : get
2 F 5 Z NEnNnNnN = ny are in the middle to higher end of divergence val-
o8| w| O A 2 ss = = x =< a ues reported for reptile (and other vertebrate) con-
Y in geners with respect to cyt b sequence divergence
5S & 2 (Seis (Johns and Avise 1998). A higher order molecular
= Sees
= . ie Sess lis phylogeny of Pachydactylus (Lamb and Bauer
i's | Si ersh Ss S's 2002) reveals that P. parascutatus is the sister spe-
| Aas Ses TS SS :
= B 2 | < S 8 sas = =°S cies of P. scutatus and that these two species (and
iver meme eee ee SR Pipe, = bly P lensis, for which ti
Sere ee Pee presumably P. angolensis, for which tissues are
Se ea reaig Yet Tate aera aero
mT) Al] anmntnoran unavailable) are members of a monophyletic group

that also includes the northern Namibian endemics

P. oreophilous and P. gaiasensis.
BIOGEOGRAPHY. — The removal of Pachydactylus robertsi from the P. scutatus group elimi-
nates the need to explain the enigmatic disjunction of more than 700 km that previously characterized
the geographic distribution of this group. The P. scutatus group as recognized here is endemic to the
region of northwestern Namibia and southwestern Angola, an area first identified as a center of reptile

34 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES
Volume 53, No. 3

endemism by Steyn and Mitchell (1965) based on the distribution of the skink Mabuya laevis. Bauer et
al. (2001) regarded this species and numerous others (e.g., Pedioplanis benguellensis, Pachydactylus
caraculicus, P. fitzsimonsi, Sepsina alberti, Hemirhagerrhis viperina, Lycophidion hellmichi,
Coluber zebrinus, Prosymna visseri) as elements of a significant trans-Kunene mopaneveld area of
endemism. Pachydactylus scutatus as presently defined is largely restricted to the numerous koppies
scattered across the northern Namibian mopaneveld. Pachydactylus parascutatus appears to be dis-
tributed on the western periphery of the mopaneveld and into the largely treeless areas of the northern
Pronamib. This distribution, as currently understood, is similar to that of the geckos Pachydactylus
oreophilus and Rhoptropus biporosus. The recognition of a new species in this region further empha-
sizes the significance of northwestern Namibia as a center of reptile diversity (Simmons et al. 1998;
Griffin 2000) and highlights the degree to which members of the genus Pachydactylus have radiated
in the western arid zone of southern Africa (Bauer 1999[2000)]).

ACKNOWLEDGMENTS

Permission to collect in the Republic of Namibia and the Skeleton Coast Park was provided by the
Ministry of the Environment and Tourism. In particular, we thank Mike Griffin for his continued sup-
port of our research in Namibia. We also thank the authorities of the Northern Cape Province and De
Beers Consolidated Farms for permission to collect Pachydactylus weberi. This research was funded
by a grant to the authors from the National Science Foundation of the United States (DEB-9707568).
Robert Drewes and Jens Vindum (CAS), Mathilda Awases (SMW), Linda Ford (AMNH), Wulf
Haacke (TM), and James Hanken and José Rosado (MCZ) kindly loaned material in their care. Randy
Babb, Paul Moler, Tori Schneider, Steve Lochetto, and Amy Meeker provided field assistance in
Namibia.

LITERATURE CITED

Amos, B. AND A. R. HOELZEL. 1991. Long-term preservation of whale skin for DNA analysis. Rep. Int. Whal.
Comm. Spec. Iss. 13:99—103.

Bauer, A. M. 1999 [2000]. Evolutionary scenarios in the Pachydactylus-group geckos of southern Africa: new

hypotheses. Afr. J. Herpetol. 48:53—62.

BAUER, A. M. AND W.R. BRANCH. 1995. Geographic variation in the Pachydactylus punctatus complex

(Reptilia: Gekkonidae) in northern Namibia. Trop. Zool. 8:69-84.

Bauer, A. M., W. R. BRANCH, AND W. D. HAACKE. 1993. The herpetofauna of the Kamanjab area and adjacent

Damaraland, Namibia. Madoqua 18:117—145.

Bauer, A. M., T. LAMB, W. R. BRANCH, AND R. D. BABB. 2001. New records of two rare snakes from northern
Namibia, with comments on the trans-Kunene distribution of mopaneveld squamates. Herpetozoa
14:75-79.

BAUER, A. M. AND A. P. RUSSELL. 1992. The evolutionary significance of regional integumentary loss in island
geckos: a complement to caudal autotomy. Ethol. Ecol. Evol. 4:343-358.

Bauer, A. M., A. P. RUSSELL, AND R. E. SHADWICK. 1989. Mechanical properties and morphological correlates
of fragile skin in gekkonid lizards. J. Exper. Biol. 145:79—102.

. 1993. Skin mechanics and morphology of two species of Pachydactylus (Reptilia: Gekkonidae). S. Afr.
J. Zool. 28:192—197.

BRANCH, W. R. 1998. Field guide to the snakes and other reptiles of southern Africa, 3rd ed. Struik Publishers,
Cape Town.

BRANCH, W.R., A. M. BAUER, AND D. A. Goop. 1996. A review of the Namaqua gecko, Pachydactylus
namaquensis (Reptilia: Gekkonidae) from southern Africa, with the description of two new species. S. Afr.
J. Zool. 31:53-69.

BROADLEY, D. G. 1977. A review of the northeastern forms of the Pachydactylus capensis complex (Sauria:
Gekkonidae). Arnoldia (Rhodesia) 8(18):1—19.

BAUER, LAMB, AND BRANCH: PACHYDACTYLUS SCUTATUS 35

FITZSIMONS, V. F. 1938. Transvaal Museum expedition to South-West Africa and Little Namaqualand, May to
August 1937. Reptiles and amphibians. Ann. Transvaal Mus. 19: 153-209 + map.

. 1943. The lizards of South Africa. Mem. Transvaal Mus. (1): i-xv + 1-528, + 24 pls. + map.

. 1959. Some new reptiles from southern A frica and southern Angola. Ann. Transvaal Mus. 23:403—-409.

Gigss, W. 1971. A preliminary vegetation map of South West Africa. Dinteria 4:5—114.

GRIFFIN, M. 2000. The species diversity, distribution and conservation of Namibian reptiles: a review. J. Namibia
Wiss. Ges. 48:116—141.

HAACKE, W. D. 1970. New herpetological records from South West Africa. Ann. Transvaal Mus. 26:277—283.

HASEGAWA, M., H. KISHINO, AND T. YANO. 1985. Dating the human-ape splitting by a molecular clock of mito-
chondrial DNA. J. Mol. Evol. 22:160—174.

HEwItT, J. 1927. Further descriptions of reptiles and batrachians from South Africa. Rec. Albany Mus. 3:71-415,
pls. 20-24.

IRISH, J. 1994. The biomes of Namibia, as determined by objective categorisation. Nav. Nas. Mus. Bloemfontein
10:549-592.

JOHNS, G. C. AND J. C. AVISE. 1998. A comparative summary of genetic distances in the vertebrates from the m1-
tochondrial cytochrome 6 gene. Mol. Biol. Evol. 15:1481—1490.

JURGENS, N. 1991. A new approach to the Namib region. I: Phytogeographic subdivision. Vegetatio 97:21—38.

KLuGE, A. G. 1993. Gekkonoid lizard taxonomy. International Gecko Society, San Diego.

. 2001. Gekkotan lizard taxonomy. Hamadryad 26:1—209.

Lamp, T. AND A. M. BAUER. 2000. Systematics of the Pachydactylus rugosus complex of southern African
geckos (Squamata: Gekkonidae). Afr. Zool. 35:55—67.

. 2002, in press. Phylogenetic relationships of the large-bodied members of the African lizard genus
Pachydactylus (Reptilia: Gekkonidae). Copeia.

LAURENT, R. L. 1964. Reptiles et amphibiens de I’ Angola (troisi¢me contribution). Comp. Diamantes Angola
Publ. Cult. 67:1—165.

LeEvITON, A. E., R. H. GIBBS, JR., E. HEAL, AND C. E. DAWSON. 1985. Standards in herpetology and ichthyology:
Part 1, Standard symbolic codes for institutional resource collections in herpetology and ichthyology.
Copeia 1985:802-832.

LOVERIDGE, A. 1944. New geckos of the genera Afroedura, new genus, and Pachydactylus from Angola. Amer.
Mus. Novit. (1254):1-4.

. 1947. Revision of the African lizards of the Family Gekkonidae. Bull. Mus. Comp. Zool. 98:1—469, pls.
1-7.

MCLACHLAN, G. R. AND J. M. SPENCE. 1966. The genus Pachydactylus (Part 1). Ann. Cape Prov. Mus.
5:149-156.

. 1967. A new species of Pachydactylus from Sesfontein, South West Africa. Cimbebasia (21):1—8.

MERTENS, R. 1955. Die Amphibien und Reptilien Siidwestafrikas, aus den Ergebnissen einer im Jahre 1952
ausgefuhrten Reise. Abhandl. Senckenberg. naturf. Ges. (490):1—172.

Meyer, A., T. D. KOCHER, P. BASASIBWAKI, AND A. C. WILSON. 1990. Monophyletic origin of Lake Victoria
cichlid fishes suggested by mitochondrial DNA sequences. Nature 347:550—-553.

ROSLER, H. 2000. Kommentierte Liste der rezent, subrezent und fossil bekannten Geckotaxa (Reptilia:
Gekkonomorpha). Gekkota 2:28—153.

SIMMONS, R. E., M. GRIFFIN, R. E. GRIFFIN, E. MARAIS, AND H. KOLBERG. 1998. Endemism in Namibia: pat-
terns, processes and predictions. Biodiv. Conserv. 7:513—530.

STEYN, W. AND A. J. L. MITCHELL. 1965. A new scincid genus: and a new record from South West Africa.
Cimbebasia (12):2—12.

. 1967. Two new gekkos from South West Africa. Cimbebasia (21):9-21.

THOMPSON, J. D., T. J. GIBSON, F. PLEWNIAK, F. JEANMOUGIN, AND D. G. HIGGINS. 1997. The CLUSTAL X win-
dows interface: flexible strategies for multiple sequence alignment aided by quality analysis tools. Nucleic
Acids Res. 24:4876-4882.

VAN DEN ELZEN, P. 1983. Zur Herpetofauna des Brandberges, Siidwest-A frika. Bonn. zool. Beitr. 34:293—309.

WERMUTH, H. 1965. Liste der rezenten Amphibien und Reptilien, Gekkonidae, Pygopodidae, Xantusiidae. Das
Tierreich 80. Walter de Gruyter, Berlin.

36 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES
Volume 53, No. 3

SPECIMENS EXAMINED

Museum acronymys follow Leviton et al. (1985). Specimens for which molecular data were ob-
tained are marked with an asterisk (*).

Pachydactylus angolensis: Angola: AMNH R-47874 (holotype), MCZ 49856 (paratype),
Hanha.

Pachydactylus parascutatus: Namibia, Kunene Region: CAS 214735* (paratype), SMW R
9454* (formerly CAS 214755; holotype), vic. Para Camp, 2 km N Sesfontein, Opuwo District
(19°07'28"S, 13°35'29”E); CAS 214749*, Purros Rd., ca. 27 km W of Sesfontein, Opuwo District
(19°03'16"S, 13°30'08"E); CAS 214750* (paratype), Purros Rd., ca. 8 km W Sesfontein, Opuwo
District (19°10'05"S, 13°34'03”"E); SMW R 182 (paratype), Kaoko Otavi, Opuwo District (1815Bc);
SMW R 184 (paratype), 20 km W of Orupembe, Opuwo District (1812Ab).

Pachydactylus robertsi: Namibia, Karas District: TM 17854 (holotype), Farm Kraikluft, Great
Karas Mountains, Great Namaqualand.

Pachydactylus sansteyni: Namibia, Kunene District: CAS 214589*, Skeleton Coast National
Park, ca. | km S of Huab River Bridge, Khorixas District (20°54'03"S, 13°32'01"E); CAS 214767*,
Skeleton Coast National Park, N bank of Huab River at Huab River Bridge, Khorixas District
(20°54'04"S, 13°31'30"E); SMW R 1623-5, 1637 (paratypes), 1626 (holotype), Kuidas (2013 Da);
SMW R 7690, 20 mi. N of Ugab river Mouth on Coast, Khorixas District.

Pachydactylus scutatus: Namibia, Kunene Region: CAS 176081, Farm Franken, Outjo District;
CAS 176112, Torrabaai Rd., 70.9km W of Kamanjab, Khorixas District; CAS 193672-3*,
Kamanjab-Torrabaai Rd., Khorixas District (19°37'27"S, 14°43'19"E); CAS 193733, Otjivize,
Opuwo-Okangwati Rd., Opuwo District (17°37'11"S, 13°27'32"”E); AMB 5943*,CAS 206961,
Sesfontein Rd., 25.3 km N Palmwag, Khorixas District; AMB 5945, Sesfontein Rd., 52 km N
Palmwag, Khorixas District (19°27'28"S, 13°52'41”E); SMW R 181, Numas Valley, Brandberg;
SMW R 183, Kamanjab, Outjo District; SMW R 185 Brandberg (2114Ba); SMW R 5183, Ugab River
atrd. 76 (20°52’S, 14°57'E);. SMW R 5722, Palmwag (19°47'S, 13°55'E); SMW R 7972, 8002, 8032,
Otjongoro, Omaruru District (20°53’S, 15°38'E).

Pachydactylus weberi: South Africa, Northern Cape Province: CAS 206754*, South Africa,
Northern Cape Province, Kleinsee, Nature Reserve (29°38'49"S, 17°05'02"E); Namibia, Erongo Re-
gion: TM 33650, Horebis Nord, Karibib District; Namibia, Khomas Region: TM 32834, 32838,
Windhoek; TM 28781, Klein Windhoek.

: _
te re Te re
pep =

lane

-_—

€ —
oe |

PROCEEDINGS OF THE CALIFORNIA ACAT

Vid Ce

Volume 53, No. 4, pp. 37-49, 3 figs.

New and Reconsidered

Thomas F. Daniel
Department of Botany, California Academy of Sciences
Golden Gate Park, San Francisco, California 94118

Three species of Justicia are newly described from the lowland rain forests of southeastern
Mexico (Oaxaca and Veracruz): J. ardens, J. tuxtlensis, and J. uxpanapensis.
Macromorphological and palynological features of these species are considered in an at-
tempt to elucidate their taxonomic affinities. New combinations in Justicia are proposed
for Beloperone nelsonii (from Nayarit) and Jacobinia stellata (from Jalisco). A new name,
Justicia veracruzana, is proposed for the species previously treated as Jacobinia paniculata
(Puebla and Veracruz).

RESUMEN

Se describen tres especies nuevas de Justicia de las selvas tropicales de la tierra baja del
sudeste de México (Oaxaca y Veracruz): J. ardens, J. tuxtlensis, y J. uxpanapensis. Se
consideran las caracteristicas macromorfoldgicas y palinologicas de estas especies para
intentar aclarar sus afinidades taxonémicas. Se proponen nuevas combinaciones en
Justicia para Beloperone nelsonii (de Nayarit) y Jacobinia stellata (de Jalisco). Un nuevo
nombre, Justicia veracruzana, se propone para la especie tratada previamente como
Jacobinia paniculata (de Puebla y Veracruz).

Justicia L. is the largest genus of Mexican Acanthaceae with about 80 species in the country
(Daniel 1999a). In addition to those species already described, there remain numerous undescribed
species, species described in genera now treated as synonymous with Justicia but that have not been
transferred to that genus, plants of uncertain taxonomic status, and names that have not been placed.
Efforts to deal with some of these “problematic” plants and names have resulted in three new species
from the lowland rain forests of Veracruz and Oaxaca, new combinations in Justicia for Jacobinia
stellata (from Jalisco) and Beloperone nelsonii (from Islas Marias, Nayarit), and a new name for
Jacobinia paniculata (from Veracruz and Puebla).

NEW SPECIES

The lowland rain forests on the Gulf slope of southeastern Veracruz and adjacent regions of
Oaxaca harbor an impressive assemblage of wet-forest Acanthaceae (Daniel 1991, 1999b, 2001).
Herewith, three additional distinctive species of Justicia are described from these forests. As has been
demonstrated elsewhere (e.g., Acosta C. and Daniel 1993; Daniel 1999b; Daniel and Wasshausen
1990; Durkee and McDade 1996), placement of new taxa from the Neotropics into the infrageneric
classification proposed by Graham (1988) is often unsatisfactory.

38 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES
Volume 53, No. 4

Justicia tuxtlensis T. F. Daniel, sp. nov.
Fig. | (a-d)

TyPE. — MEXICO. Veracruz: Mpio. San Andrés Tuxtla, Estacion de Biologia Tropical “Los
Tuxtlas,” vereda vigia I, lat. 18°34—36'N, long. 95°04—09’W, 400-500 m, 19 May 1989, G. barra M.
& S. Sinaca C. 3372 (holotype: MEXU!; isotype: CAS!).

Herbae usque ad 4 dm. Folia petiolata, laminae ovatae vel ovato-ellipticae, 36—110 mm longae,
11-43 mm latae, 2—2.8-plo longiores qua latiores. Inflorescentia floribus in spicas vel paniculas
spicarum; dichasia alterna, sessilia, uniflora. Bracteae aliquantum heteromorphae, oppositae (vel
suboppositae), lineares vel lanci-lineares, |—1.9 mm longae, 0.2—0.3 mm latae. Calyx 5-lobus,
33.7 mm longus, lobis homomorphis. Corolla alba et labio inferiore purpurato notato, 6.5—7 mm
longa, extus pubescens trichomatibus eglandulosis. Stamina thecis 0.6—1 mm longis, superpositis,
theca supera pubescens trichomatibus eglandulosis, theca inferna basi calcarata. Capsula 6—9.5 mm
longa, pubescens trichomatibus eglandulosis.

Herbs to 4 dm. Young stems subquadrate to quadrate-sulcate to + flattened, 4 grooves/lines pres-
ent, bifariously (or at least trichomes concentrated in 2 lines) pubescent with retrorse eglandular tri-
chomes 0.2—0.5 mm long, trichomes usually with prominent maroon septae. Leaves petiolate,
petioles to 22 mm long, blades ovate to ovate-elliptic, 36-110 mm long, 1 1—43 mm wide, 2—3.3 times
longer than wide, acute to acuminate at apex, acute to subattenuate at base, surfaces pubescent with
antrorse eglandular trichomes (mostly restricted to midvein) to nearly glabrous, margin entire to
subsinuate. Inflorescence of axillary and terminal dichasiate pedunculate spikes or panicles of spikes
to 9 cm long (including peduncle and excluding flowers), spikes |—2.5 mm in diameter and panicles
8-37 mm in diameter near midpoint of fertile portion, peduncles 4-15 mm long, (bifariously to)
evenly pubescent with an understory of erect to flexuose to antrorse to retrorse eglandular trichomes
0.05—0.2 mm long and an overstory of erect to flexuose glandular trichomes 0.1—0.4 mm long (glan-
dular pubescent), rachis glandular pubescent, inflorescence bracts (if present) sometimes
subfoliaceous, linear to linear-elliptic to lanceolate to lance-ovate, 2-10 mm long, 0.5—3 mm wide,
abaxial surface pubescent with antrorse eglandular trichomes, inflorescence branches (if present)
sometimes clustered (up to 5) at inflorescence nodes; dichasia alternate, sometimes + secund,
1-flowered, | per axil, sessile. Bracts opposite (to subopposite), linear to lance-linear, 1—2.2 mm long,
0.20.3 mm wide, somewhat heteromorphic with fertile bract 1.3—2 times longer than sterile bract,
abaxial surface glandular pubescent (or sometimes with glandular trichomes sparse or absent). Brac-
teoles linear to lance-linear, 1.3—2 mm long, 0.2—0.4 mm wide, abaxial surface pubescent like bracts.
Flowers sessile to subsessile (i.e., with pedicels to 0.3 mm long). Calyx 5-lobed, 3—3.8 mm long,
lobes homomorphic, lance-subulate, 2.4-3 mm long, 0.4—0.6 mm wide, abaxially pubescent like
bracts, margins + hyaline. Corolla whitish with maroon markings on lower lip, 6.5—8 mm long, exter-
nally pubescent with flexuose to retrorse eglandular trichomes 0.1—0.2 mm long, tube + expanded dis-
tally, 4-5 mm long, 1.2—1.4 mm in diameter near midpoint, upper lip 2—3 mm long, 2-lobed, lobes
(.1—0.2 mm long, lower lip 2-3 mm long, lobes 0.8—1.3 mm long, I|—1.6 mm wide. Stamens inserted
near apex of corolla tube (less than 1 mm from mouth of corolla), 2.5—3 mm long, filaments glabrous
or with one or more coarse trichomelike protrusions distally, thecae 0.6—1 mm long, subequal in
length, parallel, superposed (gap 0.2—0.3 mm long), distal theca pubescent with flexuose eglandular
trichomes, lower theca with a prominent pointed basal appendage 0.2—0.4 mm long; pollen (Fig. 2)
3-colporate, 6-pseudocolpate, exine between colpi and pseudocolpi sometimes appearing as if break-
ing up into insulae. Style 4.5—-5.5 mm long, proximally pubescent with eglandular trichomes, distally
glabrous, stigma 0.1—0.2 mm long, asymmetric, lobes inconspicuous. Capsule 6—9.5 mm long, pu-
bescent with erect to flexuose eglandular trichomes 0.05—0.2 mm long (inconspicuous sessile glands

DANIEL: ACANTHACEAE IX. JUSTICIA Bo.

FIGURE 1. a-d, Justicia tuxtlensis (Ibarra M. & Sinaca C. 3372).a. habit, x 0.5. b. anthers, x 10. c. capsule, x 4. d. seed, x 6. e-i,
J. ardens. e. leaf (Wendt et al. 4227), x 0.4. f. inflorescence (Wendt et al. 4227), x 0.7. g. anthers (Wendt et al. 4227), x 4.5. h.
capsule (Wendt et al. 4765), x 2.3. i. seed (Croat & Hannon 63269), x 3.8. j-o, J. uxpanapensis. j. habit (Vazquez et al. 839), x
0.5. k. fertile bract (Vazquez et al. 839), x 3.1. sterile bract (Vazquez et al. 839), x 3. m. anthers (Wendt et al. 3328), x 9. n. cap-
sule (Poole et al. 1493), x 3.5. 0. seed (Poole et al. 1493), x 6. Scale for a, | = 15 mm; for b = 0.8 mm; for c = 1.9 mm; for

d=1.3mm; for e= 19mm; for f=11 mm; for g=1.6mm; for h=3.2 mm; for i=2 mm; for k, 1=2.5 mm; for m,
o = 1.2 mm; forn=2.1 mm.

40 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES
Volume 53, No. 4

usually evident near apex as well), stipe 2-4 mm long, head subellipsoid (often with a slight medial
constriction), 4-5.5 mm long. Seeds 4, lenticular, 1.5—2.2 mm long, 1.5—2 mm wide, surfaces and
margin covered with low moundlike protrusions, lacking trichomes.

PHENOLOGY. — Flowering and fruiting: April-May.

DISTRIBUTION AND HABITAT. — Southeastern Mexico (Veracruz); plants occur in lowland rain
forest (“selva alta perennifolia”) at 400-500 m elevation.

PARATYPES. — MEXICO. Veracruz: Estacion de Biologia Tropical Los Tuxtlas, Cerro Vigia, A.
Gentry et al. 32250 (US); between Catamaco and Zontecomapan on road to Monte Pio, ca. 7 mi. from
Catamaco, H. Moore & M. Cetto 6254 (US); Mpio. San Andres Tuxtla, Est. Biol. Trop. “Los
Tuxtlas,” Cerro del Vigia, 7. Ramamoorthy 2321 (EB).

In the corolla tube (well below the point of insertion of the stamens) there are two prominent tri-
angular invaginations along the veins to the upper lip. These appear to be homologous with (or identi-
cal to) the staminodelike thickenings encountered in some other species with heteromorphic bracts
such as Justicia chol T. F. Daniel, J. costaricana Leonard, and J. nevlingii Wassh. & T. F. Daniel
(Wasshausen and Daniel 1995; see Fig. 1, f), as well as J. uxpanapensis T. F. Daniel (see below). Gra-
ham (1988:559) noted that among species from both the Old World and the New World, “the bases of
the filaments and of the rugula and the veins beneath them are usually pubescent and may also be pro-
duced into the corolla tube as pouch-like processes. These may almost occlude the tube . . .” Hedrén
(1989) noted the occurrence of similar invaginations among homomorphically bracted species in the
Paleotropical section Harnieria (Solms-Laub.) Benth. He also noted these invaginations help to close
the access route to the nectar disc and young fruit for small insects.

Justicia tuxtlensis shows similarities to several tropical American species including J. metallica
Lindau and J. pectoralis Jacq. of sect. Sarotheca (Nees) Benth. (Graham 1988) and J. comata (L.)
Lam. This latter species was treated as a taxon of uncertain infrageneric affinities by Graham (1988).
These four species can be distinguished using the key below. Additionally, J. pectoralis (Mexico,
Central America, West Indies, and South America to Bolivia) differs from J. tuxtlensis by its
unifarious cauline pubescence; subulate bracts and bracteoles; equally to subequally inserted, gla-
brous, and basally unappendaged thecae; 2-aperturate pollen; and capsules with glandular pubes-
cence. Justicia metallica (Costa Rica and Panama) further differs from Justicia tuxtlensis by its pink
to purplish corollas, subperpendicular to perpendicular and glabrous thecae, 2-aperturate pollen, and
echinate seed surfaces. In several morphologically conspicuous characters (e.g., clusters of spikes at
some inflorescence nodes, calyx form, and corolla size) Justicia tuxtlensis is similar to the widespread
(Mexico, Central America, West Indies, and South America to Argentina) J. comata. It can be readily
distinguished from that species by the characters noted in the key.

1. Calyx 5-lobed, the lobes homomorphic; corolla 3—7 mm long.

2. Inflorescence rachises and abaxial surfaces of bracts, bracteoles, and calyx lobes glabrous or pubescent with
eglandular trichomes only; corolla 3—6 mm long; thecae 0.3—0.5 mm long, glabrous, lacking basal appendages;
pollenZzapenunatescapsules—4 minis On lane enon cures Utne ee J. comata

2. Inflorescence rachises and abaxial surfaces of bracts, bracteoles, and calyx lobes pubescent with glandular and
eglandular trichomes; corolla 6.5—8 mm long; thecae 0.6—1 mm long, the upper theca pubescent with eglandular
trichomes, the lower theca with a basal appendage 0.2—0.4 mm long; pollen 3-aperturate; capsule 6—9.5 mm long.

Se ORS SR AM POR inc Shenae ey iti, Ro eT Mae OIE OLE? Eee do eo 0 J. tuxtlensis
1. Calyx 4-lobed with the lobes homomorphic or 5-lobed with the lobes heteromorphic (i.e., posterior lobe greatly reduced
in size relative to the others); corolla 7.5—14 mm long.

3. Young stems unifariously pubescent; corolla 7.5—10 mm long; calyx 2—3.5 mm long; thecae subparallel to
parallels meave/zeky se ey eaiyta ee eee Peccten ANG to tae Tera baa keg cg on fey a alae ea a J. pectoralis

3. Young stems bifariously pubescent; corolla 12—14.5 mm long; calyx 47.5 mm long; thecae subperpendicular to
PEUPENGICUIAL sed ee a) nyt Pa, eee Oe ase ney Sceae trie rd ee aneion hae a J. metallica

DANIEL: ACANTHACEAE IX. JUSTICIA 4]

FIGURE 2. SEM images of pollen. a. Justicia tuxtlensis (Ibarra M. & Sinaca C. 3372), apertural view. b. J. tuxtlensis (Ibarra
M. & Sinaca C. 3372), interapertural view. c. J. ardens (Wendt et al. 2822), apertural view. d. J. ardens (Wendt et al. 4227),
interapertural view. e. J. uxpanapensis (Wendt et al. 3328), apertural view. f. J. uxpanapensis, (Wendt et al. 3328), subpolar
view. Scale for a = 26 um, for b and e = 32 um, for c and d = 44 um, for f= 41 um.

The epithet of this species is derived from the name of the region in which it appears to be en-
demic.

Justicia ardens T. F. Daniel, sp. nov.
Fig. 1 (e-1).

TyPE. — MEXICO. Veracruz: Mpio. Minatitlan, Cerro Blanco, ca. 7 km NE de Uxpanapa (Pob.
12) en camino a Pob. 15, ca. 17°14’N, 94°09’W, 200 m, 19 October 1983, T. Wendt et al. 4227
(holotype: CAS!; isotypes: to be distributed from CHAPA).

42 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES
Volume 53, No. 4

Frutices usque ad 2.5 m alti. Folia petiolata, laminae ovato-ellipticae vel ellipticae, 105-290 mm
longae, 49-128 mm latae, 1.6—3.3-plo longiores quam latiores. Inflorescentia floribus in fasciculos
terminales densos; dichasia + opposita, sessilia vel subsessilia, uniflora. Bracteae homomorphae, +
oppositae, oblanceolatae, 7-11 mm longae, 1.5—2.5 mm latae. Calyx 5-lobus, 8—11 mm longus, lobis
homomorphis. Corolla aurantiaca et apice lutea, 50-60 mm longa, extus pubescens trichomatibus
glandulosis. Stamina thecis 2.2—3.2 mm longis, impariter insertis vel superpositis, basi ecalcaratis,
glabris vel theca supera pubescens trichomatibus eglandulosis. Capsula 12.5—17 mm longa, glabra.

Shrubs to 2.5 m tall. Young stems quadrate to quadrate-sulcate, bifariously pubescent with
antrorse eglandular trichomes 0.2—0.5 mm long. Leaves petiolate, petioles to 50 mm long, blades
ovate-elliptic to elliptic, 105-290 mm long, 49-128 mm wide, 1.6—3.3 times longer than wide,
acuminate to abruptly short-acuminate at apex, acute to attenuate at base, abaxial surface glabrous,
adaxial surface pubescent with antrorse eglandular trichomes along midvein, margin entire to + irreg-
ularly subsinuate. Inflorescence of axillary (in axils of distalmost pair of leaves) and terminal
pedunculate panicles of modified spikes collectively forming a dense terminal subcapitate cluster of
flowers, individual panicles 15—32 mm long (including peduncle and excluding flowers), fertile por-
tion 10-20 mm in diameter near midpanicle, peduncles of panicles to 6 mm long, rachises and
peduncles pubescent like young stems; dichasia + opposite, 1-flowered, 1 per axil, sessile to
subsessile. Bracts + opposite, oblanceolate, 7-11 mm long, 1.5—2.5 mm wide, abaxially glabrous,
margin ciliate with flexuose eglandular trichomes to | mm long. Bracteoles oblanceolate to linear,
6-11 mm long, 0.7—1.5 mm wide, abaxial surface glabrous, margin ciliate like bracts. Flowers ses-
sile. Calyx 5-lobed, 8—11 mm long, lobes lanceolate, subequal, 7—9.5 mm long, 0.9—1.2 mm wide,
abaxially glabrous, margin ciliate with flexuose eglandular trichomes. Corolla subfusiform in bud,
orangish with yellow at apex, 50-60 mm long, externally pubescent with erect to flexuose glandular
trichomes 0.05—0.6 mm long, tube gradually expanded from near base, 30-40 mm long, 34.2 mm in
diameter near midpoint, upper lip 19-22 mm long, 2-lobed, lobes 0.5—0.6 mm long, lower lip
18-22 mm long, 3-lobed, lobes 2.5-4 mm long, |—1.5 mm wide. Stamens inserted in distal 1/3 of co-
rolla tube, 26-30 mm long, filaments glabrous distally, sparsely pubescent with eglandular trichomes
near base, thecae 2.2-3.2 mm long, equal to subequal (distal theca often longer), + parallel, unequally
inserted (overlapping by up to 0.9 mm) to superposed (contiguous), glabrous (or distal theca apically
pubescent with a few eglandular trichomes), lacking basal appendages; pollen (Fig. 2) 3-colporate,
6-pseudocolpate, exine between colpi and pseudocolpi sometimes breaking up into insulae. Style
48-58 mm long, glabrous, stigma subequally to unequally 2-lobed, lobes 0.05—0.2 mm long. Capsule
12.5-17 mm long, glabrous, stipe 6.5—9 mm long, head subglobose, 6—8 mm long. Seeds 4 (or fewer
by abortion), lenticular, subcordate, 3.5—3.8 mm long, 3—3.5 mm wide, immature surfaces and mar-
gin covered with slender trichomelike protrusions less than 0.5 mm long, mature surfaces and margin
smooth, glabrous.

PHENOLOGY. — Flowering: October-November; fruiting: December—January.

DISTRIBUTION AND HABITAT. — Southeastern Mexico (Oaxaca, Veracruz); plants occur in low-
land rain forests (with Terminalia, Dialium, Guatteria, Ficus, Pouteria, Poulsenia, Symphonia,
Sterculia, Guarea, Crataevia) at 100-350 m elevation.

PARATYPES. — MEXICO. Oaxaca: Uxpanapa Region, along road between Esmeralda (17 km E
of Sarabia) and Rio Manea, | 1.5—13.5 mi S of Esmeralda, 17°04’N, 94°45’ W, T. Croat & D. Hannon
63269 (CAS); Mpio. Sta. Maria Chimalapa, Congregacion Nicolas Bravo, ca. 3-5 km S de la pob. Rio
Alegre, Veracruz, Arroyo de la Cascada, cerca del Rancho Rutt, ca. 17°10’N, 94°42°W, T. Wendt et
al. 4765 (CAS). Veracruz: Mpio. Hidalgotitlan, reserva ejidal, laderas del Cerro Amarillo, ca.
17°47°N, 94°39°W, G. Castillo C. 348 (F); Mpio. Minatitlan, lomas S del Poblado 11 y al S de la
brecha 105, ca. 27 km E de La Laguna, ca. 17°14’N, 94°18’W, 7. Wendt et al. 2822 (CAS).

DANIEL: ACANTHACEAE IX. JUSTICIA 43

The spikes of this species are unusual in that the bracts and bracteoles are rotated 90° from their
normal position and they are fused basally forming a short pedunclelike stalk. The flowering inflores-
cences are usually so dense that these peculiarities are not readily evident.

The infrageneric affinities of this species are not obvious using Graham’s (1988) infrageneric
classification. Justicia ardens shares several diagnostic characters (e.g., compound spicate inflores-
cences, long and yellowish corollas) with species (e.g., J. aurea Schltdl.) in section Cyrtanthera
(Nees) V.A.W. Graham. Pollen grains among species in that section, however, are described as
2-aperturate with three to four rows of insulae on each side of the apertures. Among species of Justicia
with yellow or orange corollas more than 50 mm long in the Uxpanapa-Chimalapa regions of
Veracruz and Oaxaca, J. ardens can be distinguished using the following key:

1. Inflorescences subtended by cupulate involucres formed by 2 bracts connate for (1/4—) 1/3—1/2 their length, fused proxi-
mal portion 15—35 mm long; calyx 15—25 mm long; corolla externally glabrous; thecae 5—7.3 mm long; capsule
EOP mT ONOMSECUS S11 OMT ONG amare mere Rabe ey Nel eesieene. cosa! Gl Zecliel | often ae ee J. borrerae
1. Inflorescences not subtended by cupulate involucres as above; calyx 3.5—11 mm long; corolla externally pubescent;
thecae 1.9-3.5 mm long; capsule 10-23 mm long, seeds 34 mm long.
2. Corolla externally pubescent with eglandular trichomes only, trichomes yellowish; calyx lobes 1.5—2.8 mm wide;
Ehecae wlth conspicious basalkappendares=m teeth seh c es cl mene) se echscy Sys SES See eee J. fimbriata
2. Corolla externally pubescent with glandular (and sometimes also eglandular) trichomes, trichomes not yellowish;
calyx lobes 0.9-1.3 mm wide; thecae lacking basal appendages.
3. Inflorescence of elongate panicles; bracts abaxially pubescent; calyx 3.5—-8 mm long, abaxially pubescent;
corolla yellow, externally pubescent with glandular and eglandular trichomes, lobes of lower lip 1-2 mm
long, buds cobralike (i.e., erect but prominently curved near apex so that distal portion of bud is horizontal);
capsule pubescentsced papillatepolleni2-apertuiates = aes) 2s syes0s) 4 4) oes e cle een aaemnenee J. aurea
3. Inflorescence of dense terminal subcapitate clusters; bracts abaxially glabrous; calyx 8-11 mm long,
abaxially glabrous; corolla orangish with yellow at apex, externally pubescent with glandular trichomes only,
lobes of lower lip 2.5—4 mm long, buds erect throughout (i.e., not cobralike); capsule glabrous; mature seed
Smoothamollenks-apentttatese erty ate ai en: ciksat cases 's oiea-cy so) hace ecto omar Gh Oe RCN J. ardens

The epithet of this species, meaning burning or glowing in Latin, refers to the dense clusters of
brightly colored flowers that must appear like flames in the forest understory.

Justicia uxpanapensis T. F. Daniel, sp. nov.
Fig. 1 (j-o).

Type. — MEXICO. Veracruz: Mpio. Hidalgotitlan, brecha La Laguna-E] Elefante, 17°17'N,
94°30’ W, 150 m, 26 July 1974, M. Vazquez et al. 839 (holotype: MEXU!; isotypes: MO! NCU!).

Herbae perennes usque ad 2.5 dm altae. Folia petiolata, laminae ovatae vel ellipticae vel anguste
ellipticae, 29-80 mm longae, 12—32 mm latae, 2.2-4.6-plo longiores quam latiores. Inflorescentia
floribus in spicas; dichasia alterna, sessilia, uniflora. Bracteae heteromorphae, alternae; bracteae
fertiles ellipticae vel late ellipticae, 1 1—14 mm longae, 5—8.5 mm latae; bracteae steriles lanceolatae
vel ellipticae, 7-11 mm longae, 1.5—3.5 mm latae. Calyx 5-lobus, 5—6 mm _ longus, lobis
homomorphis. Corolla rosea vel purpurea, 14.5—-18 mm longa, extus pubescens trichomatibus
eglandulosis. Stamina thecis 1-1.4mm longis, impariter insertis, theca supera pubescens
trichomatibus eglandulosis, theca inferna basi calcarata. Capsula 9-10 mm longa, pubescens
trichomatibus eglandulosis.

Erect to spreading perennial herbs to 2.5 dm tall. Young stems quadrate-sulcate, + bifariously pu-
bescent with an understory of antrorsely appressed eglandular trichomes 0.2—0.5 mm long and +
evenly pubescent with an overstory of flexuose eglandular trichomes 1—3 mm long. Leaves petiolate,
petioles to 7 mm long, blades ovate to elliptic to narrowly elliptic, 29-79 mm long, 12-30 mm wide,

44 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES
Volume 53, No. 4

2.24.6 times longer than wide, acute to acuminate at apex, acute to subattenuate at base, surfaces pu-
bescent with flexuose to antrorse eglandular trichomes, margin entire. Inflorescence of axillary and
terminal pedunculate dichasiate spikes to 75 mm long (including peduncle and excluding flowers),
7—17 mm in diameter near midspike, peduncles 34.5 mm long, pubescent like young stems, rachis +
evenly pubescent with antrorse to antrorsely appressed eglandular trichomes 0.2—0.6 mm long; dicha-
sia alternate, secund, sessile, | per axil, 1-flowered. Bracts sometimes drying dark, alternate,
heteromorphic, fertile bracts larger than sterile bracts, petiolate, elliptic to broadly elliptic, 11-14 mm
long, 5—8.5 mm wide, abaxial surface and margin pubescent with flexuose eglandular trichomes
12.2 mm long, sterile bracts petiolate, lanceolate to elliptic, often asymmetric, 7-11 mm long,
1.5-3.5 mm wide, pubescent like fertile bracts. Bracteoles linear, 6-10 mm long, 0.4—2 mm wide, pu-
bescent like bracts. Flowers sessile. Calyx 5-lobed, 5-6 mm long, lobes homomorphic, lanceolate,
4.5—5 mm long, 0.4—0.9 mm wide, abaxially very sparsely pubescent with antrorse eglandular tri-
chomes along midvein 0.1—0.6 mm long, margin and apex with a few antrorse eglandular trichomes.
Corolla pinkish or purplish, 14.5-18 mm long, externally pubescent with flexuose eglandular tri-
chomes 0.2—0.7 mm long, tube 9—9.5 mm long, gradually expanded distally, 3 mm wide near mid-
point, upper lip 5.5—7 mm long, 2-lobed, lobes 0.6—0.8 mm long, lower lip 6—8.5 mm long, lobes
3-5 mm long, 2.5—6 mm wide. Stamens inserted near apex of corolla tube, 5—5.7 mm long, filaments
glabrous, thecae 1—1.4 mm long, subequal in size (lower larger), subparallel to subperpendicular, un-
equally inserted (overlapping by 0.2—0.4 mm), upper theca pubescent with eglandular trichomes,
lower theca with a basal appendage 0.3—0.4 mm long; pollen (Fig. 2) 2-aperturate, apertures flanked
on each side by | (—2) rows of insulae. Style 9.3 mm long, glabrous, stigma subcapitate to minutely
and + equally 2-lobed, 0.1 mm long. Capsule 9-10 mm long, distally pubescent with flexuose
eglandular trichomes 0.2—0.3 mm long, stipe 3.5—4 mm long, head subellipsoid with a slight medial
constriction, 5—6.3 mm long. Seeds 4 per capsule, sublenticular, 2—2.8 mm long, 1.4—2.1 mm wide,
surfaces sometimes + wrinkled and forming ridges, surfaces and margin covered with low moundlike
protuberances.

PHENOLOGY. — Flowering and fruiting: May—July.

DISTRIBUTION AND HABITAT. — Southeastern Mexico (Veracruz); plants occur in lowland rain
forests at 150—170 m elevation.

PARATYPES. — MEXICO. Veracruz: brecha, near Poblado 2, Uxpanapa military reserve, J.
Poole et al. 1493 (MEXU, TEX); Mpio. Hidalgotitlan, lomitas al SE del Poblado 6, 17°16’N,
94°30'W, T. Wendt et al. 3328 (CAS).

As in several other heteromorphically bracted species (see above under J. tuxtlensis), the internal
surface of the corolla tube (below midpoint) has 2 prominent invaginations, each pubescent with
eglandular trichomes. Pollen of this species (based on Wendt et al. 3328) has a furrowlike polar con-
nection between trema regions. I have not observed similar furrows in pollen of other Mexican
Justicia (Daniel 1998).

Using Graham’s (1988) infrageneric classification, J. uxpanapensis appears somewhat similar to
species in sect. Simonisia (Nees) V.A.W. Graham. Unlike species in that section, however, in J.
uxpanapensis the fertile bracts are elliptic to broadly elliptic (vs. triangular, lanceolate, or
oblanceolate), the corollas are 14.5—18 mm long (vs. 19-64 mm long), pollen grains have 24 rows of
insulae in the trema regions (vs. 4-6 rows in similar grains of sect. Simonisia), and the seeds are
sublenticular (vs. not compressed) and glabrous (vs. pubescent). Affinities of this species are proba-
bly best sought among other heteromorphically bracted species from southern Mexico such as J.
neviingii and J. chol. These morphologically similar species can be distinguished using the following
key:

DANIEL: ACANTHACEAE IX. JUSTICIA 45

1. Corolla pinkish or purplish, 14.5-18 mm long; bracts alternate, sterile bracts 7-10 mm long; calyx 5—6 mm long; pollen
Jpapeninmics Crue Qa) mMN NN, se aoe ease gocoo no sb no wee hoo c BE aa AB Oe J. uxpanapensis
1. Corolla white to yellow or greenish yellow with maroon markings, 8.5—11.5 mm long; bracts opposite, sterile bracts
2-6 mm long; calyx 2.5—5 mm long; pollen 3—4-aperturate; capsule 6—8.5 mm long.
2. Young stems with trichomes concentrated in 2 lines; inflorescence of axillary dichasiate spikes; bracteoles
0.4-1 mm wide; style pubescent; seeds 4, covered with low rounded papillae or ridges; pollen 3-aperturate. J. chol
2. Young stems with trichomes restricted to 4 lines; inflorescence of (axillary and) terminal panicles of dichasiate
spikes; bracteoles 1.2—2.2 mm wide; style glabrous; seeds 2, smooth; pollen 4-aperturate........ J. neviingii

NEW COMBINATIONS AND A NEW NAME

Beloperone Nees and Jacobinia Nees are now usually treated as congeneric with Justicia (e.g.,
Graham 1988). Although nomenclatural renovations for most taxa described in these genera have
been published, names do not appear to exist in Justicia for several Mexican taxa. Two new combina-
tions and a new name are proposed below for three of them. Complete descriptions that permit com-
parison of these taxa with other species of Justicia are also provided.

Justicia stellata (B. L. Rob. & Greenm.) T. F. Daniel, comb. nov.

Jacobinia stellata B. L. Rob. & Greenm., Proc. Amer. Acad. Arts 29:390. 1894. Type. — MEXICO. Jalisco: dry
cliffs, barranca of Tequila, 16 Oct 1893, C. Pringle 4573 (holotype: GH!; isotypes: MEXU, NSW!, NY!,
US!).

Perennial herbs or shrubs of unknown height. Young stems subterete to subquadrate, densely and
evenly pubescent with cream-colored stellate to dendritic eglandular trichomes 0.2—1 mm long.
Leaves petiolate, petioles to 20 mm long, blades ovate to ovate-elliptic, (22—) 30-135 mm long, (6—)
9-64 mm wide, 2.3—3.4 (—3.7) times longer than wide, acute to acuminate at apex, acute to
subattenuate at base, surfaces pubescent with stellate to dendritic trichomes, the abaxial surface usu-
ally more densely so, margin entire to subcrenate. Inflorescences of axillary and/or terminal
pedunculate dichasiate thyrses to 60 mm long (including peduncle and excluding flowers), ca. 20 mm
in diameter near midpoint of fertile portion, thyrses collectively forming a terminal panicle, axillary
thyrses alternate or opposite, | per axil, pedunculate, peduncles to 22 mm long, pubescent like young
stems and with erect glandular trichomes 0.2—0.5 mm long as well, rachises pubescent like peduncles
(except with eglandular trichomes sparser); dichasia alternate, 1-flowered, often + secund, | per axil,
pedunculate, peduncles 2-4 mm long, pubescent like rachis. Bracts sessile to petiolate, opposite, lin-
ear to narrowly elliptic to obovate-elliptic, 7.5-17 mm long, 1.7—-4 mm wide, abaxial surface pubes-
cent like rachis. Bracteoles linear to oblanceolate-spatulate, 6.5-10.5 mm long, 1—1.8 mm wide,
abaxial surface pubescent like rachis. Flowers sessile to subsessile. Calyx 5-lobed, 17-21 mm long,
lobes linear to oblanceolate-spatulate, subequal, 15—18 mm long, 1—2.2 mm wide, abaxially pubes-
cent like rachis. Corolla “reddish purple” (fide protologue), 36-44 mm long, externally pubescent
with glandular (and a few unbranched eglandular trichomes) 0.2—0.4 mm long, tube gradually ex-
panded distally, 21-26 mm long, 2.8-3 mm in diameter near midpoint, upper lip 18-19 mm long,
2-lobed, lobes 0.5—0.6 mm long, lower lip 17-19 mm long, spreading, lobes 3 mm long, 3-3.5 mm
wide. Stamens inserted near apex of corolla tube, 18-19 mm long, filaments distally pubescent with
glandular trichomes 0.05—0.1 mm long, thecae 2.5-2.9mm_ long, equal to subequal,
subperpendicular, unequally inserted (overlapping by 2 mm), pubescent (especially upper theca) with
glandular trichomes 0.05—0.1 mm long, lower theca with a spurlike basal appendage to 0.2 mm long;
pollen (Fig. 3) 2-aperturate, apertures flanked on each side by 2 rows of insulae. Style 35-42 mm long,
glabrous, stigma equally 2-lobed, lobes 0.2 mm long. Capsule 13—15 mm long, pubescent with glan-
dular trichomes 0.1—0.2 mm long, stipe length unknown, head shape and length unknown. Seed num-

46 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES
Volume 53, No. 4

ber and shape unknown, 3.5 mm long, 3 mm wide, surfaces papillose, papillae pubescent with
trichomes less than 0.05 mm long.

PHENOLOGY. — Flowering and fruiting: October.

DISTRIBUTION AND HABITAT. — Western Mexico (Jalisco); although the precise habitat remains
unknown, the type locality is in a region of tropical deciduous forest and thornscrub at an elevation
less than 1200 m.

The presence of branched trichomes is unusual among American Acanthaceae and their abun-
dance in this species makes it readily identifiable.

I am not aware that this species has ever been collected subsequent to Pringle’s collection in
1893. It is odd that a plant with such large and colorful flowers would not have been recollected in
more than 100 years in a region that has been visited by numerous botanists. The species must be e1-
ther very rare or no longer extant.

Justicia nelsonii (Greenm.) T. F. Daniel, comb. nov.

Beloperone nelsonii Greenm., Proc. Amer. Acad. Arts 33: 488. 1898. TypE. — MEXICO. Nayarit: Islas Marias,
Maria Madre Island, 3-25 May 1897, E. Nelson 4246 (holotype: GH!; isotype: US!).

Perennial herbs to 5 dm tall. Young stems subquadrate, + evenly (or trichomes + concentrated in
2 lines) and sometimes densely pubescent with erect to flexuose eglandular trichomes 0.2—0.7
(—1) mm long. Leaves petiolate, petioles to 25 mm long, blades ovate to elliptic, 30-90 mm long,
11-40 mm wide, 2.4—2.8 times longer than wide, acute at apex, acute to subattenuate at base, surfaces
pubescent with erect to flexuose to antrorsely appressed eglandular trichomes, margin entire to
subsinuate-crenate. Inflorescence of terminal pedunculate + densely bracteate dichasiate spikes to
40 mm long (including peduncles and excluding flowers), 12—14 mm in diameter near midpoint of
fertile portion, peduncles to 6 mm long, pubescent like young stems, rachis evenly pubescent with
erect to flexuose eglandular trichomes 0.2—0.6 mm long; dichasia opposite, sessile, 1 per axil,
1-flowered. Bracts opposite, obovate to elliptic, narrowed proximally, 8-11 mm long, 2.8—5 mm
wide, apically erect, abaxial surface pubescent with erect to flexuose to antrorse eglandular (to
subglandular) and sometimes glandular trichomes 0.05—0.5 mm long. Bracteoles linear-elliptic to lin-
ear to lance-linear, 6—9 mm long, 1.1—2.1 mm wide, abaxial surface pubescent like bracts. Flowers
sessile. Calyx 5-lobed, 4.5—5.5 mm long, lobes homomorphic, lanceolate to lance-subulate,
3.34.3 mm long, 0.7—0.9 mm wide, abaxially pubescent like bracts. Corolla white, 21—23 mm long,
externally pubescent with flexuose eglandular trichomes 0.2—0.5 mm long, tube 11—13 mm long,
2.8-3.3 mm in diameter near midpoint, gradually expanded distally, upper lip 9—9.7 mm long,
2-lobed, lobes 0.8 mm long, lower lip 10—11.5 mm long, lobes 3.5—5 mm long, 1.6—2.6 mm wide.
Stamens 9-10 mm long, inserted near apex of corolla tube, filaments pubescent with eglandular tri-
chomes, thecae 1.3—1.8mm long, subequal, subparallel, unequally inserted (overlapping by
0.5—1 mm), dorsally pubescent with flexuose eglandular trichomes, each with a bulbous basal ap-
pendage to 0.4 mm long (appendage of lower theca larger than that of upper theca); pollen (Fig. 3) 3
(—4)-aperturate (see Discussion). Style 14-19 mm long, pubescent with upward-pointing eglandular
trichomes, stigma 0.2 mm long, lobes unequal 0.1—0.2 mm long. Capsule 9—11 mm long, pubescent
with flexuose eglandular trichomes 0.1—0.2 mm long, head subellipsoid, 6.5—7 mm long. Seeds 4,
3.2 mm long, 2.5 mm wide, surfaces minutely papillose, margin with inconspicuous apically barbed
trichomelike protuberances (at least on portions of the margin).

PHENOLOGY. — Flowering: May, October; fruiting: May.

DISTRIBUTION AND HABITAT. — Pacific islands off the west coast of Mexico (Nayarit); plants
occur on canyon floors at an unknown elevation.

DANIEL: ACANTHACEAE IX. JUSTICIA 47

FiGurE 3. SEM images of pollen. a. Justicia stellata (Pringle 4573), apertural view. b. J. nelsonii (Nelson 4246), apertural
veiw. c. J. nelsonii (Nelson 4246), interapertural view. d. J. nelsonii (Nelson 4246), polar view. e. J. veracruzana (Chazaro & H.
de Chazaro 4078), apertural view. f. J. veracruzana (Chazaro & H. de Chazaro 4078), interapertural view. Scale for a and
e = 32 um, for b and d= 19 um, for c = 23 um, for f = 35 um.

ADDITIONAL SPECIMENS EXAMINED. — MEXICO. Nayarit: Tres Marias Islands, Maria Madre
Island, R. Ferris 5692 (DS), F. Maltby 137 (US).

Considerable variation in pubescence is evident among the few known collections of this species.
Maltby 137 and the isotype at US have glands on the bracts, bracteoles and calyx. These are not evi-
dent on Ferris 5692 or the holotype. The young stems of Ferris 5692 are also more densely pubescent
than those of the other collections.

In the protologue of Beloperone nelsonii, Greenman (1898) noted that this species closely resem-
bles B. comosa Nees (= Justicia fulvicoma Schltdl. & Cham.) but has a shorter corolla with a broader
lower lip and somewhat larger leaves with longer and less pubescent petioles. These taxa are indeed
quite similar. Comparison of both species reveals that they can be distinguished by the following cou-
plet:

48 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES
Volume 53, No. 4

Corolla white, 21-23 mm long; bracts 2.8—5 mm wide, apically erect; stamens 9-10 mm long; style 14-19 mm long.

Per etek Mare eng em ae ann Gs Vi tic ota hes 1h GRY Ene eR A A POR epee ha OE CM SS 5 J. nelsonii
Corolla orange, 25-38 mm long; bracts 6-12 mm wide, apically recurved-spreading; stamens 13—17 mm long; style
30=34 mimVlonee cage a sor Che eo ee a ot Bh urstetins ce eee ee Cee 2 eh oe J. fulvicoma

Some abnormalities were noted in pollen of Nelson 4246 (Fig. 3) including unusual twisting.
Whereas most grains appeared to be 3-aperturate, some were 4-aperturate. On each side of the aper-
tures, there is some evidence of the exine separating to form indistinct insulae.

Justicia veracruzana T. F. Daniel, nom. nov.

Jacobinia paniculata Oerst., Vidensk. Meddel. Dansk Naturhist. Foren. Kjobenhavn 1854:153. 1855, not
Justicia paniculata of earlier authors [e.g., N. L. Burm. (1768), Forssk. (1775), Rose (1895), Sessé & Moc.
(1887)]. Type. — MEXICO. Veracruz: Colipa, March 1841, F. Liebmann s.n. (10674) (syntypes: C!,
CAS!, P!, US!); Mirador, March 1842, F. Liebmann s.n. (10675) (syntype: C!).

Perennial herbs to 2 m tall. Young stems terete to subquadrate, + bifariously to + evenly pubes-
cent with flexuose antrorse to antrorsely appressed eglandular trichomes 0.1—0.3 mm long, trichomes
with conspicuous maroon septae, soon glabrate. Leaves petiolate, petioles to 40 mm long, blades
ovate-elliptic to elliptic to obovate, 35-180 mm long, 21—85 mm wide, |.8—2.5 times longer than
wide, often asymmetric, round to acute to subacuminate at apex, acute to subattenuate at base, sur-
faces sparsely pubescent with antrorse to antrorsely appressed eglandular trichomes, trichomes + re-
stricted to major veins, margin entire to crenate. Inflorescence of pedunculate dichasiate spikes (to
racemes) or panicles of spikes (to racemes) from axils of distal leaves and inflorescence bracts, collec-
tively forming an open terminal panicle to 18 cm long, 4—7.5 cm in diameter near midpoint of fertile
portion, peduncles of spikes (to racemes) or panicles to 30 mm long, pubescent like young stems; in-
florescence bracts opposite, often caducous, linear to linear-elliptic, 4-6 mm long, 0.8—1 mm wide,
abaxial surface pubescent with antrorsely appressed eglandular trichomes; spikes (to racemes) or pan-
icles opposite, |—2 per axil, rachis pubescent like young stems; dichasia alternate, often + secund,
1-flowered, | per axil, sessile. Bracts often caducous, triangular to subulate, 1—1.5 mm long, 0.5 mm
wide, abaxial surface pubescent with antrorse to antrorsely appressed eglandular trichomes
0.05—0.1 mm long. Bracteoles often caducous, triangular to linear, sometimes conduplicate,
1.22 mm long, 0.3—0.6 mm wide, abaxial surface pubescent like bracts. Flowers subsessile to
pedicellate, pedicels 0.5—1.5 mm long. Calyx 5-lobed, 9.5-18 mm long, lobes lanceolate to
lance-linear to linear-elliptic (to oblanceolate), equal to subequal, 8-16 mm long, 1.5—3.5 mm wide,
abaxially sparsely pubescent (at least proximally) with antrorsely appressed eglandular trichomes
0.05—0.1 mm long. Corolla yellow, 31—36 mm long, externally pubescent with erect to flexuose glan-
dular and eglandular trichomes 0.1—0.3 mm long, tube 14-17 mm long, 3 mm in diameter near mid-
point, + gradually expanded distally, throat indistinct, upper lip 16-18 mm long, 2-lobed, lobes
0.4 mm long, lower lip 15—17 mm long, lobes 2.5-4.5 mm long, 1—1.5 mm wide. Stamens inserted
near apex of corolla tube, 17—19 mm long, thecae subparallel to subperpendicular, 2.5—3.3 mm long,
subequal, unequally inserted (overlapping by 1.5—2.2 mm), glabrous, lower theca with a + bulbous
basal appendage 0.2—0.3 mm long; pollen (Fig. 3) 2-aperturate, multiechinate. Style 29-34 mm long,
distally glabrous, stigma subcapitate, 0.1—0.2 mm long, lobes inconspicuous. Capsule 14-17 mm
long, glabrous, stipe S—7 mm long, head ellipsoid to obovoid, 9.5—10.5 mm long. Seeds 4, lenticular,
23.5 mm long, 2.5—3.2 mm wide, surfaces and margin tuberculate with rounded to subconic protu-
berances bearing minute projections.

PHENOLOGY. — Flowering: January—March, October; fruiting: January—March, October.

DISTRIBUTION AND HABITAT. — Southeastern Mexico (Puebla, Veracruz); plants occur in tropi-
cal deciduous forest at 1650 m elevation.

DANIEL: ACANTHACEAE IX. JUSTICIA 49

ADDITIONAL SPECIMENS EXAMINED. — MEXICO. Guanajuato: cultivated in Guanajuato,
1889, A. Duges 75,c (US). Puebla: Mpio. Patlanalan, camino del paraje El Aguacate, Ver. a
Patlanalan, Pue., M. Chazaro & P. Hernandez de Chazaro 4078 (US). Veracruz: Zacuapan, Barranca
de Tenampa, C. Purpus 4334 (US); Barranca del Fortin San Martin Tlacatopec, C. Purpus 16780
(US).

Most of the collections of J. veracruzana were made more than 70 years ago. However, Chazaro
& Hernandez de Chazaro 4078, collected in 1986, reveals the relatively recent persistence of this spe-
cies in nature. Pollen of J. veracruzana conforms to Graham’s (1988) “Type 9,” which lacks a trema
area and is covered with short, subconical echinae. Such pollen was noted by Graham (1988) in sev-
eral species of her sect. Plagiacanthus (Nees) V.A.W. Graham. Justicia veracruzana differs from
Graham’s (1988) description of that section by it seeds that are tuberculate (vs. smooth) and lack a dif-
ferentiated margin (vs. “with a distinct margin’).

ACKNOWLEDGMENTS

I am grateful to Jenny Speckles for preparing the line drawing, Scott Serata for assisting with the
scanning electron microscope, Gary Williams and Darrell Ubick for help with the digital images of
pollen, Fernando Chiang and Roberto Gonzalez T. for checking herbaria at MEXU and IBUG for
specimens of Justicia stellata, and the curators of the following herbaria for making specimens avail-
able: C, CAS, DS, F, GH, IEB, MEXU, MO, NCU, NY, P, TEX, and US.

LITERATURE CITED

AcosTA C., S. AND T. F. DANIEL. 1993. A new species of Justicia (Acanthaceae) from southern Mexico. Kew
Bull. 48:119—123.

DANIEL, T. F. 1991. A revision of Aphelandra (Acanthaceae) in Mexico. Proc. Calif. Acad. Sci. 47:235—274.

. 1998. Pollen morphology of Mexican Acanthaceae: diversity and systematic significance. Proc. Calif.

Acad. Sci. 50:217—256.

. 1999a. Acanthaceae. Fasc. 23, pp. 1-102 in Flora del Valle de Tehuacan-Cuicatlan, P. D. Davila A. et

al., eds. Univ. Nac. Auton. México.

. 1999b. Taxonomic and distributional notes on Neotropical Justicia (Acanthaceae). Proc. Calif. Acad.

Sci. 51:483—492.

. 2001. Streblacanthus monospermus (Acanthaceae), a genus and species new to the flora of Mexico.
Contr. Univ. Michigan Herb. 23:139—144.

DANIEL, T. F. AND D. C. WASSHAUSEN. 1990. Three new species of Justicia (Acanthaceae) from Panama. Proc.
Calif. Acad. Sci. 46:289-297.

DurKEE, L. H. AND L. A. MCDADE. 1996. Three new species of Justicia (Acanthaceae) from Costa Rica. Novon
6:13-21.

GRAHAM, V. A. W. 1988. Delimitation and infra-generic classification of Justicia (Acanthaceae). Kew Bull.
43:551—-624.

GREENMAN, J. M. 1898. Diagnoses of new and critical Mexican Phanerogams. Proc. Amer. Acad. Arts
33:471-489.

HEDREN, M. 1989. Justicia sect. Harnieria (Acanthaceae) in tropical Africa. Acta Univ. Ups. Symb. Bot. Ups.
29:1-141.

WASSHAUSEN, D. C. AND T. F. DANIEL. 1995. Justicia nevlingii (Acanthaceae), a new species from Mexico.
Novon 5:114—-117.

= ¥ 7 = ; 8s a :
be New Species 678 7 | ;
Beawiiendeida Fad
SbF bs! (ic

PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES

Volume 53, No. 5, pp. 51—61, 8 figs. June 14, 2002

| JUN 80 2002

Sponging off of Porifera: New Species of Cryptic Dorid
Nudibranchs (Mollusca, Nudibranchia) from the Tropical
Indo-Pacific

Terrence M. Gosliner
Department of Invertebrate Zoology and Geology, California Academy of Sciences
Golden Gate Park, San Francisco, California 94118

and

Angel Valdés
Natural History Museum of Los Angeles County
900 Exposition Boulevard, Los Angeles, California 90007

Two new species of Asteronotus are described from the Indo-Pacific tropics. Asteronotus
mimeticus sp. nov. is described from specimens collected in the Philippines, Palau, Austra-
lia, and Papua New Guinea. Asteronotus spongicolus sp. nov. is described from the three lo-
calities in Tanzania and one in northern Queensland, Australia. Both species are
remarkably cryptic on their prey sponges. Asteronotus mimeticus is variable in its color and
notal ornamentation, depending upon the sponge on which it is feeding. Both species differ
from the only recognized species in the genus, A. cespitosus (van Hasselt, 1824) in their
smaller body size, lack of regular ridges and rows of tubercles and differences in the
radular morphology and reproductive system. In A. cespitosus the outermost radular tooth
is elongate while in the two new species the outermost tooth is reduced to an ovoid plate.
The accessory gland of A. cespitosus is regular in shape with a straight stylet while in the
two new species the accessory gland is irregularly shaped with a curved stylet. Asteronotus
mimeticus has a rounded body shape while A. spongicolus is more elongate and ovoid. In A.
mimeticus, the inner radular teeth are usually devoid of denticles, but may rarely have 1—2
additional denticles on the outer side of the inner 14 radular teeth. In A. spongicolus the in-
nermost tooth has 1—2 denticles on its inner side and the succeeding 7—11 teeth have 1—2
denticles on the outer side of the tooth. The outermost teeth of A. mimeticus are rounded
ovoid plates while those of A. spongicolus are more quadrangular with an acute apex. The
reproductive system of A. mimeticus has a bursa copulatrix that is smaller than the
receptaculum seminis while in A. spongicolus the receptaculum is larger than the bursa.
The genital atrium of A. mimeticus is shorter and narrower than that of A. spongicolus. The
accessory spine of A. mimeticus has a narrow base and is gradually curved while in A.
spongicolus the spine has a broad base with a sharply curved apex.

Recent field studies at several localities around the Indo-Pacific tropics have yielded numerous
individuals of dorid nudibranchs that are cryptic on large, foliose sponges inhabiting shallow water
coral reef communities. Anatomical studies indicate that these dorids represent two undescribed spe-
cies of the genus Asteronotus Ehrenberg, 1831. The only currently recognized member of this genus is
A. cespitosus (van Hasselt, 1824). Valdés and Gosliner (2001) have recently reviewed its anatomy and
systematic position. The present paper describes the anatomy of these two new species. Type material

Nn
i)

PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES
Volume 53, No. 5

is housed in the collections of the California Academy of Sciences (CASIZ) and the Natural History
Museum of Los Angeles County (LACM).

SPECIES DESCRIPTIONS

Asteronotus mimeticus sp. nov.
Figs. LA=D; 2; 3; 4; 5; 6A, B

MATERIAL EXAMINED. — HOLOTYPE: CASIZ 157487, Arthur’s Rock, Calumpan Peninsula,
Batangas Province, Luzon Island, Philippines, 15 m depth, 22 February 1992, T. M. Gosliner.
PARATYPES: CASIZ 084294, six specimens, same locality and date as the holotype. LACM 2901, one
specimen, same locality and date as the holotype. CASIZ 156622, three specimens, two dissected, Ar-
thur’s Rock, Calumpan Peninsula, Batangas Province, Luzon Island, Philippines, 15 m depth, 11 May
2001, T. M. Gosliner. CASIZ 156664, one specimen, Devil’s Point, Maricaban Island, Batangas
Province, Luzon Island, Philippines, 15 m depth, 10 May 2001, T. M. Gosliner. CASIZ 157492, two
specimens, one dissected, Layag Layag, Caban Island, Batangas Province, Luzon Island, Philippines,
15 mdepth, 7 May 2001, T. M. Gosliner. CASIZ 157491, one specimen, Layag Layag, Caban Island,
Batangas Province, Luzon Island, Philippines, 8 m depth, 7 May 2001, A. Valdés. CASIZ 083709,
four specimens, Koala, Calumpan Peninsula, Batangas Province, Luzon Island, Philippines, 28 Feb-
ruary 1992, T. M. Gosliner. CASIZ 088127, five specimens, Twin Rocks, Calumpan Peninsula,
Batangas Province, Luzon Island, Philippines, 17 m depth, 22 March 1993, T. M. Gosliner. CASIZ
156594, one specimen, Koala, Calumpan Peninsula, Batangas Province, Luzon Island, Philippines,
15 m depth, 12 May 2001, T. M. Gosliner. CASIZ 088155, two specimens, one dissected, 2 km e. of
Lighthouse, Dakak, Mindanao Island, Philippines, 20 m depth, 29 March 1993, T. M. Gosliner.
CASIZ 086032, one specimen, N. side of Lighthouse, Dakak, Mindanao Island, Philippines, 31
March 1993, T. M. Gosliner. CASIZ 113637, one specimen, dissected, Sullivan’s Patches, n. side of
Milne Bay, Milne Bay Province, Papua New Guinea, 27 May 1998, T. M. Gosliner. CASIZ 113693,
three specimens, one dissected, Tagula Island, Calvados Chain, Louisiade Archipelago, Papua New
Guinea, 4 June 1998, T. M. Gosliner. CASIZ 113695, one specimen, Brooker Channel, Calvados
Chain, Louisiade Archipelago, Papua New Guinea, | June 1998, T. M. Gosliner. CASIZ 109732, two
specimens, one dissected, inner side of barrier reef, Babalukes, Koror Municipality, Palau, 23 Sep-
tember 1996, T. M. Gosliner. CASIZ 139593, one specimen, Jemeluk Beach, Amed, Bali, Indonesia,
3 m depth, 31 October, 2000, T. M. Gosliner. OTHER MATERIAL: Australian Museum, AM C115955,
one specimen, 21 mm alive, Lizard Island, North Queensland, Australia, 100 m off research lab on
patch reef, 3 m depth, on Carteriospongia sp., 1 June 1979, J. Bakus and W. B. Rudman.

ETYMOLOGY. — The name Asteronotus mimeticus is derived from its resemblance to its sponge
prey with which it is cryptically associated.

NATURAL HISTORY. — This species is found abundantly on the undersides of the leafy sponge,
Phyllospongia lamellosa (Esper, 1799) and a few other foliose demosponges such as Carteriospongia
sp., where the nudibranchs are cryptically colored on these species of sponges (Fig. 1A—D). The egg
mass of this species (Fig. 2) is a flat, tightly coiled spiral of 2-3 whorls. The eggs are relatively large,
suggesting that this species may have direct or lecithotrophic rather than plaktotrophic development.

DISTRIBUTION. — This species is known from several localities in the western Pacific Ocean
from the Philippine Islands, Palau, Indonesia, Australia (W. B. Rudman, personal communication)
and Papua New Guinea (present study).

EXTERNAL MORPHOLOGY. — The living animals (Figs. | A—D, 2) are oval in shape, 13-30 mm
in length. The body color is gray, brown or yellow with scattered opaque white spots on the notum.
The notum may be entirely smooth (Figs. 1A, B; 2), or may possess tubercles and a few elongate
papillae (Fig. 1C, D). The rhinophores are perfoliate with 6-10 lamellae. There may be brownish pig-

Nn
o>)

GOSLINER AnD VALDES: CRYPTIC DORID NUDIBRANCHS

FicurE 1. Photographs of living animals. A. Asteronotus mimeticus, brown smooth specimen from Luzon, Philippines
(CASIZ 156664); B. Asteronotus mimeticus, gray smooth specimen from Mindanao, Philippines (CASIZ 088155); C.
Asteronotus mimeticus, papillated specimen from Luzon, Philippines (CASIZ 084294), D. Asteronotus mimeticus,
tuberculated specimen from Luzon, Philippines (CASIZ 156622); E. Asteronotus spongicolus, specimen from northern Tanza-
nia (CASIZ 099293); F. Asteronotus spongicolus, specimen from southern Tanzania (CASIZ 099294). Photographs by T. M.
Gosliner.

ment on the base and middle region: the apex is opaque white. The two rhinophoral sheaths have 7-8
lobes along their margin. The gill consists of 5—6 tripinnate branchial leaves, which are the same color
as the body and may bear scattered opaque white spots. The gill pocket contains six distinct lobes. The
foot is broad and generally the same coloras the rest of the body. Anteriorly it is bilabiate and notched.
Anelongate, digitiform oral tentacle is present laterally on either side of the labial region and mouth.
LABIAL CUTICLE AND RADULA. — The buccal mass is muscular throughout. The anterior end of
the buccal mass contains the labial cuticle. which is thin throughout its length and is devoid of any
thickening, armature or rodlets. The radula is wide with numerous hamate teeth. The radular formula
is 51 x 28.0.28 (CASIZ 156622), 42 x 26.0.26 (CASIZ 156623), 48 x 32.0.32 (CASIZ 084294) and
28 x 32.0.32 (CASIZ 113693) in four specimens examined. The inner lateral teeth (Figs. 3A, B: 4A,

54 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES
Volume 53, No. 5

FIGURE 2. Asteronotus mimeticus from the Philippines with the egg mass, photograph by A. Valdés.

C) are relatively short with a broad base and a curved primary cusp. These teeth lack denticles with the
exception of a specimen from Papua New Guinea (Fig. 4C) (CASIZ 113693) where the inner 14 teeth
bear 1—2 small, triangular denticles along the outer side of the cusp. There is an abrupt transition be-
tween the inner and middle lateral teeth. The middle lateral teeth (Fig. 3C) are larger and more elon-
gate than the inner ones and lack denticles in all specimens examined. The outer lateral teeth are
elongate and curved (Figs. 3D; 4B, D) with the exception of the outermost tooth, which is reduced to
an ovoid plate.

REPRODUCTIVE SYSTEM. — The hermaphroditic reproductive system is triaulic (Fig. 5). The
ampulla is elongate and slightly convoluted. It narrows and divides into the vas deferens and oviduct.
The vas deferens expands into a short, wide prostate that consists of two distinct portions. The prostate
abruptly narrows distally into an elongate, muscular, ejaculatory portion. This muscular segment ex-
pands as it enters the relatively short common genital atrium. An irregular accessory gland is present
and has a muscular duct that enters the atrium between the deferent duct and the vagina. Within the
muscular base of the accessory gland there is an elongate, somewhat-curved cuticular spine (Fig. 6A,
B). The vaginal duct is thin and elongate and enters the large, spherical bursa copulatrix. Adjacent to

GOSLINER AND VALDES: CRYPTIC DORID NUDIBRANCHS

FIGURE 3. Scanning electron micrographs of the radula of Asteronotus mimeticus. A. Innermost lateral teeth of a smooth
specimen from the Philippines (CASIZ 156623); B. Innermost lateral teeth of a tuberculated specimen from the Philippines
(CASIZ 156622); C. Mid-lateral teeth of a tuberculated specimen from the Philippines (CASIZ 156622); D. Outermost teeth of
a tuberculated specimen from the Philippines (CASIZ 156622).

PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES
Volume 53, No. 5

FiGuRE 4. Scanning electron micrographs of the radula of Asteronotus mimeticus. A. Innermost lateral teeth of a papillated
specimen from the Philippines (CASIZ 084294): B. Outermost teeth of a papillated specimen from the Philippines (CASIZ
084294): C. Innermost lateral teeth of a smooth specimen from Papua New Guinea (CASIZ 113693); D. Outermost teeth of a
smooth specimen from Papua New Guinea (CASIZ 113693).

GOSLINER AND VALDES: CRYPTIC DORID NUDIBRANCHS 57

the proximal end of the vagina and also emerg-
ing from the bursa is another elongate duct that
joins the bursa with the smaller, thick-walled
receptaculum seminis. Also joining this duct at
the base of the receptaculum is the short uterine
duct.

Asteronotus spongicolus n. sp.
Piss bE. 6G: 728

MATERIAL EXAMINED. — HOLOTYPE:
CASIZ 099389, one specimen, Uroa, 3 kmN. of
Uroa Beach Hotel, east coast of Zanzibar, Tan-
zania, 2m depth, 10 November 1994, T. M.
Gosliner. PARATYPES: LACM 2902, one speci-
men, same locality and date as holotype. CASIZ
099293, six specimens, three dissected,
Mbudya Island, off Kunduchi Beach, Pwani Re-
gion, N. of Dar es Salaam, Tanzania, 1—3 m
depth, 27 October 1994, T. M. Gosliner. CASIZ

; Sa 099394, two specimens, one dissected, Mana

FIGURE 5. Reproductive system of Asteronotus mimeticus : : :

(CASIZ 084294), scale bar= 1 mm. Abbreviations: a, ampulla; Huanja Island, Mtwara Region, Tanzania, | No-

ag, accessory gland; be, bursa copulatrix; dd, deferent duct; fg, vember 1994, T.M. Gosliner. OTHER

female glands; pr, prostate; sr, seminal receptacle; v, vagina. MATERIAL: Australian Museum, AM C115954,

one specimen, 28mm alive, Lizard Island,

North Queensland, Australia, 100 m offresearch lab on patch reef, 3 m depth, on Carteriospongia sp.,
1 June 1979, J. Bakus and W. B. Rudman.

ETYMOLOGY. — The name Asteronotus spongicolus is derived from its association with lobate,
foliose sponges.

NATURAL HIStory. — This species 1s found abundantly on the undersides of the leafy lobate
sponge, Carteriospongia sp., where it is cryptically colored. Asteronotus spongicolus 1s found in shal-
low water of 1—3 m depth.

DISTRIBUTION. — This species is known from three localities in Tanzania, from Zanzibar and
north of Dar es Salaam south to the Mtwara region near the Mozambique border and northern
Queensland, Australia (W. B. Rudman, personal communication).

EXTERNAL MORPHOLOGY. — The living animals (Fig. 1E,F) are elongate, ovoid in shape. Liv-
ing specimens are 10-35 mm in length. The body color is yellowish green to brown with scattered,
opaque white and dark brown to black spots on the notum. There is a medial longitudinal line of lighter
pigment that extends from immediately anterior to the rhinophores to the anterior edge of the
branchial pocket. The notum may be entirely smooth or may possess a few slightly elevated tubercles.
The rhinophores are perfoliate with 10 lamellae. Opaque white spots are present on the base and mid-
dle region of the rhinophores and the apex is also opaque white. The two rhinophoral sheaths have 8
lobes along their margin. A concentration of opaque white spots is present along the margin of either
rhinophoral sheath. The gill consists of 5—6 tripinnate branchial leaves, which are the same color as
the body and bear many scattered, opaque white spots. The gill pocket contains six distinct lobes and
also bears a concentration of opaque white spots along its margin. The foot is broad and generally the
same color as the rest of the body. Anteriorly it is bilabiate and notched. An elongate, digitiform oral
tentacle is present laterally on either side of the labial region and mouth.

58 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES
Volume 53, No. 5

10um

FIGURE 6. Scanning electron micrographs of the copulatory spines. A. Asteronotus mimeticus smooth specimen from the
Philippines (CASIZ 084294), B. Asteronotus mimeticus, tuberculated specimen from the Philippines (CASIZ 15622); C.
Asteronotus spongicolus, smooth specimen from Tanzania (CASIZ 099293).

LABIAL CUTICLE AND RADULA. — The buccal mass is muscular throughout. The anterior end of
the buccal mass contains the labial cuticle, which is thin throughout its length and is devoid of any
thickening, armature or rodlets. The radula is wide with numerous hamate teeth. The radular formula
is 43 x 30.0.30 (CASIZ 099293) and 52 x 27.0.27 (CASIZ 099394) in two specimens examined. The
inner lateral teeth (Fig. 7A,B) are relatively short with a broad base and a curved primary cusp. The in-
nermost teeth have 1—2 denticles on the inner side of the primary cusp. The succeeding 7 teeth have
1-2 small, triangular denticles on the outer side of the cusp. There is an abrupt transition between the
inner and middle lateral teeth. The middle lateral teeth (Fig. 7C) are larger and more elongate than the
inner ones and lack denticles in both specimens examined. The outer lateral teeth are elongate and
curved (Fig. 7D) with the exception of the outermost tooth, which is reduced to a quadrangular plate
with an acute anterior apex.

REPRODUCTIVE SYSTEM. — The hermaphroditic reproductive system 1s triaulic (Fig. 8). The
ampulla is thick, elongate and slightly curved. It narrows and divides into the vas deferens and ovi-
duct. The vas deferens expands into a short, wide prostate that consists of two distinct portions. The
prostate abruptly narrows distally into an elongate, muscular, ejaculatory portion. This muscular seg-
ment expands as it enters the relatively long, wide common genital atrium. An irregular accessory
gland is present and has a long convoluted muscular duct that enters the atrium between the deferent
duct and the vagina. Within the muscular base of the accessory gland is an short, strongly curved cutic-
ular spine with a broad base (Fig. 6C). The vaginal duct is thin and elongate and enters the small,
spherical bursa copulatrix. Adjacent to the proximal end of the vagina and also emerging from the
bursa is another elongate duct that joins the bursa with larger, thick-walled receptaculum seminis.
Also joining this duct at the base of the receptaculum is the short uterine duct, which then enters the fe-
male gland mass.

GOSLINER AND VALDES: CRYPTIC DORID NUDIBRANCHS

FiGuRE 7. Scanning electron micrographs of the radula of Asteronotus spongicolus (CASIZ 099293). A. Innermost lateral
teeth: B. Detail of the denticles on the innermost lateral teeth; C. Mid-lateral teeth; D. Outermost lateral teeth.

60 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES
Volume 53, No. 5

DISCUSSION

The anatomy and systematics of the dorid
nudibranch genus Asterontous has been well
studied by recent workers (Kay and Young
1970; Edmunds 1971; Valdés and Gosliner
2001). All of these authors describe the anatomy
of A. cespitosus and consider this as the only
valid species within the genus. Valdés and
Gosliner (2001) conducted a phylogentic analy-
sis of caryophyllidia-bearing dorids and con-
cluded that Asteronotus is the sister taxon of
Halgerda Bergh, 1880. Together these two taxa
are the sister group of the caryophyllidia-
bearing dorids.

Valdés and Gosliner (2001) discussed the
possible synonymy of Peronodoris Bergh,

Re 2 : os 1904, with Asteronotus. No definitive synon-
Creu Repontine sytem of Aero sonst ymy could be made owing to the lack of type
ag, accessory gland; be, bursa copulatrix; dd, deferent duct; fg, material for the type species of Peronodoris, P.
female glands; pr, prostate; sr, seminal receptacle; v, vagina. — cancellata Bergh, 1904. Since Bergh’s 1904 de-

scription, two other species, P. denticulata
Eliot, 1908 and P. rehderi Marcus and Marcus, 1970, have been assigned to this genus. These species
all have interconnecting ridges on the notum and numerous denticulate radular teeth. Peronodoris
rehderi has a large muscular accessory structure that contains a spine (Marcus and Marcus 1970).
Species of Asteronotus have a glandular accessory structure with a muscular duct that contains a
spine. This appears to represent a significant distinction between the taxa, but further clarification of
this apparent difference must await rediscovery of species of Peronodoris to confirm these anatomi-
cal features.

The genus Asteronotus is distinguished by several autapomorphies. It has a gill with a lobed
branchial pocket, a short prostatic portion of the vas deferens and an accessory gland that bears a sin-
gle spine within a muscular structure. The two new species described here share all of these character-
istics with A. cespitosus, but differ from this species in several regards. Both species differ from A.
cespitosus in their smaller body size, lack of regular ridges and rows of tubercles and differences in the
radular morphology and reproductive system. In A. cespitosus the outermost radular tooth is elongate
while in the two new species the outermost tooth is reduced to an ovoid plate. The accessory gland of
A. cespitosus is regular in shape with a straight stylet, whereas in the two new species the accessory
gland is irregularly shaped with a curved stylet. Asteronotus mimeticus has a rounded body shape
while A. spongicolus is more elongate and ovoid. In A. mimeticus, the inner radular teeth are usually
devoid of denticles, but may rarely have 1—2 additional denticles on the outer side of the inner 14
radular teeth. In.4. spongicolus the innermost tooth has 1—2 denticles on its inner side and the succeed-
ing 7-11 teeth have 1—2 denticles on the outer side of the tooth. The outermost teeth of A. mimeticus
are rounded ovoid plates, whereas those of 4. spongicolus are more quadrangular with an acute apex.
The reproductive system of A. mimeticus has a bursa copulatrix that is larger than the receptaculum
seminis, whereas in A. spongicolus the receptaculum is larger than the bursa. The genital atrium of A.
mimeticus is shorter and narrower than that of A. spongicolus. The accessory spine of A. mimeticus has
a narrow base and is gradually curved, whereas in 4. spongicolus the spine has a broad base with a
sharply curved apex.

GOSLINER AND VALDES: CRYPTIC DORID NUDIBRANCHS 61

Many species of sponge-feeding dorid nudibranchs are known to exhibit special resemblance to
their prey sponges (Gosliner and Behrens 1990). This crypsis has been best demonstrated for temper-
ate nudibranchs and has not been as well documented for tropical taxa. This paper describes crypsis
for two new species of Asteronotus. In A. mimeticus color pattern and texture vary considerably and
reflect the variability of the sponge prey species fed upon by this species. Not only are the adult nudi-
branchs of this species cryptic, but the egg mass is also cryptically colored on the prey sponge. Most
dorid nudibranchs have an egg mass morphology, in which the edge of the mass 1s attached by one
edge of the egg ribbon and the remaining portion is free and well elevated from the surface. In A.
mimeticus, the egg mass is attached directly to the sponge by its widest margin, minimizing the visibil-
ity of the egg mass. The egg mass morphology of A spongicolus remains unknown. Nevertheless, the
color pattern and texture of these two species make them remarkable examples of special resemblance
in tropical dorid nudibranchs.

ACKNOWLEDGMENTS

We thank Kit Stewart for supporting the field work undertaken in Tanzania, the Christensen Re-
search Institute for supporting work in the Madang area of Papua New Guinea and the Coral Reef Re-
search Foundation for supporting field work in Milne Bay and the Louisiade Archipelago in Papua
New Guinea and Palau. The research on this material and production of this paper was supported by
the National Science Foundation through the PEET Grant DEB-9978155 (Phylogenetic systematics
of dorid nudibranchs). Yolanda Camacho, Shireen Fahey, Mike Miller, Vic Smith, Yvonne Valles
and Gary Williams kindly assisted with field collections. Bill Rudman, kindly provided photographs
and additional records of both species from Australia.

LITERATURE CITED

BERGH, R. 1904. Malacologische Untersuchungen, Band 9. Pp. 1-55, pls 1-4 in Reisen im Archipel der
Philippinen, C. Semper, ed. C. W. Kreidel, Wiesbaden.

EDMUNDS, M. 1971. Opisthobranchiate Mollusca from Tanzania (Suborder: Doridacea). Zoological Journal of
the Linnean Society 50:339-396.

ELIOT, C. N. E. 1908. Reports on the marine biology of the Sudanese Red Sea. XI. Notes on a collection of nudi-
branchs from the Red Sea. Journal Linnean Society London, (Zoology) 31:86—122.

GOSLINER, T. M. AND D. W. BEHRENS. 1990. Special resemblance, aposematic coloration and mimicry in
opisthobranch gastropods. Pp. 127—138 in Adaptive coloration in invertebrates, M. Wicksten, ed. American
Society of Zoologists, Galveston, Texas.

Kay, E. A. AND D. K. YOUNG. 1970. The Doridacea (Opisthobranchia; Mollusca) of the Hawaiian Islands. Pa-
cific Science 23:172—231.

Marcus, ER. AND Ev. MARCUS. 1970. Opisthobranch mollusks from the southern tropical Pacific. Pacific Sci-
ence 24:155—179.

VALDES, A. AND T. M. GOSLINER. 2001. Systematics and phylogeny of the caryophyllidia-bearing dorids
(Mollusca, Nudibranchia), with the description of a new genus and four new species from Indo-Pacific deep
waters. Zoological Journal of the Linnean Society 133:103—198.

PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES

Volume 53, No. 6, pp. 63-72, 5 figs. July 1, 2002

Five New Chordeumatidan Millipeds from China:
New Species of Vieteuma (Kashmireumatidae) and Nepalelta

SSS

(Megalotylidae) | wooMatine Biolo ical Laboratory’

by
William A. Shear JUL 08 2002
Department of Biology, Hampden-Sydney College,
Hampden-Sydney, Virginia 23943
Woods Hole, WA 02544

Five new species of chordeumatidan millipeds, Vieteuma longi (Kashmireumatidae),
Nepalella pianma, N. griswoldi, N. kavanaughi and N. magna (Megalotylidae), are de-
scribed from the Gaolingong Shan, Yunnan Province, China. The new species bring to
seven the total number of chordeumatidans described from the Chinese mainland.

Our knowledge of chordeumatidan millipeds from China is very limited compared to the pro-
spective fauna of that country. No definitive record of the order from China had been published before
Mauriés and Nguyen-Duy Jaquemin (1997) recorded a single species, Vieteuma hubeiensis
(Kashmireumatidae), a troglobite from Shennonghia, Hubei Province. To put this in an entomological
perspective, it would be analogous to recording the first butterfly from China in 1997!

More recently, I described a second troglobite, Nepalella caeca (Megalotylidae), from
Shuicheng, Guizhou Province (Shear 1999a). That paper summarizes past tentative reports of
chordeumatidans from China; Wang and Mauriés (1996) have provided a valuable summary of the
myriapodology of China in general. Shear (1999b) discussed chordeumatidan millipeds from Taiwan.

In 1998, the California Academy of Sciences sponsored a collecting trip to China, and a number
of chordeumatidan millipeds were collected in Yunnan Province, in and around the Gaolingong Shan.
Charles Griswold kindly forwarded them to me for study. Included were four new species of
Nepalella, and a single new Vieteuma, all surface-dwelling forms. It might have been thought more
advisable to hold this material and wait for more substantial revisionary work on the two genera, but
given the sporadic availability in the west of collections from China, and the scanty information on
chordeumatidans there, I’ve decided to describe the new species quickly. Perhaps this work will stim-
ulate more collecting of this most interesting order of millipeds, and systematic work on them, by Chi-
nese myriapodologists.

Thanks are due the collectors listed below, and especially to Charles Griswold for bringing the
specimens to my attention. The habitat descriptions accompanying the species accounts are from
notes provided by Dr. Griswold. Specimens are deposited in the Kunming Institute of Zoology (KIZ),
Yunnan China, and California Academy of Sciences (CAS), San Francisco, USA. This study was
made possible by a grant from the National Science Foundation to the author and Petra Sierwald (NSF
DB-9712438). This is contribution 20 from the California Academy of Sciences’ Center for
Biodiversity Research and Information (CBRI) and contribution 14 from the Academy’s China Natu-
ral History Project (CNHP).

64 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES
Volume 53, No. 6

SYSTEMATIC ACCOUNT

Order Chordeumatida Koch, 1847
Suborder Craspedosomatidea Brolemann, 1935
Superfamily Brannerioidea Cook, 1896
Family Kashmireumatidae Mauriés 1982
Genus Vieteuma Golovatch 1984

This family consists of two genera, Kashmireuma Mauriés, of the Himalaya in Kashmir and Ne-
pal, and Vieteuma Golovatch, from Vietnam and southern China.

Including the following, Vieteuma contains only three species. The generotype, V. topali
Golovatch, is from Vietnam (Golovatch 1984); it is a very small species with only 26 segments in both
sexes. As mentioned above, V. hubeiensis Mauriés and Nguyen-Duy Jaquemin is a Chinese
troglobite; it has 28 segments. Vietewma females have normal second legs. The posterior gonopod
prefemora are strongly lobed, and the seventh legs of the males are slightly modified.

Kashmireuma Mauriés contains three Himalayan species, of which K. schawallari Shear and K.
nielseni Mauriés appear to be closely related, based on similarities of the anterior gonopods and the
nearly identical, strongly modified coxae of male legpair 7. Kashmireuma nepalensis Maurics has es-
sentially unmodified legpair 7 coxae and quite different gonopods (Mauriés 1988). All previously de-
scribed species of Kashmireuma have 28 segments. A unique feature among chordeumatidan
millipeds is found in this genus: the second legpair of the female is vestigial, reduced to small,
unarticulated nubbins on the second sternum. Kashmireuma males may have the posterior gonopod
prefemora extended distal to the prefemoral/femoral joint, but without the strong lobes seen in
Vieteuma.

The present species has an anterior gonopod plan which agrees well with that of V. hubeiensis,
and strongly lobed posterior gonopod coxae typical of Vieteuma. Despite this close agreement in male
characters, the females have vestigial second legs, as in Kashmireuma. Thus we must know a greater
range of species in these two genera to properly assess this female character.

Given the three species known, it is now possible to assay at least a preliminary general descrip-
tion of Vieteuma gonopods. Much of the bulk of the anterior gonopod is composed of a
synangiocoxosternum, a fusion of the sternite and the bodies of the coxae on each side. The sternum
generally has some form of mid-line extension, which in the present species takes the shape of two
large ensiform processes. The synangiocoxite (formed from the coxal walls) is cupped and bears a
subtriangular fimbriated process laterally; this process can be very large, as in V. hubeiensis. Poste-
riorly from one to three pairs of pseudoflagella are present, each with dense fimbriation. The smooth
processes emerging from the center of the angiocoxal mass are taken to be colpocoxites (formed from
the extruded and sclerotized coxal glands). The presence of articulated telopodites, seen in
kashmireumatids, is rare in chordeumatidan anterior gonopods and seems to occur most frequently in
cleidogonoids and brannerioids. The posterior gonopods have coxites that are similar in shape in the
three known species; the largest part of the gonopod is the prefemur with its thorny lobes. These thorns
can be seen under high magnification (400 x) to be short, stout setae. The reflexed prefemora are usu-
ally rather small and probably do not have a glandular function. The trochanteral modifications of
legpairs 11-16 have not been described in the other two species.

The cyphopods are unusually formed, with enlarged mesal valves. The curious, reduced, second
legpair has already been referred to; this is a character of Kashmireuma and is not found in either pre-
viously known species of Vieteuma. The entire second segment of a female V. longi (Fig. 1D) was di-
gested in trypsin, and I found a number of features not previously noted. The reduced second sternum
articulates with strong processes extending from the anterioventral margins of the pleurotergites.
There are two small, completely fused nubbins on the sternite that represent the vestigial legs. The

SHEAR: CHORDEUMATIDAN MILLIPEDS FROM CHINA 65

sternite also bears two very thin apodemes, but I could not find any sign of a spiracle. The receptacle
and valves are sparsely setose (setae are omitted for clarity in Fig. 1D) and their cuticle is scaly. Not
shown in the drawing are two small sclerites dorsal and posterior to the cyphopods and embedded in
the membrane that ties them to the second sternum; I have no ideas about the homologies of these
sclerites.

Vieteuma longi, new species
Figure |

MATERIAL EXAMINED. — Male holotype, 2 male and 4 female paratypes from China, Baoshan
Prefecture, pass over Gaolingshan at 2100 m, Nankang, 36 air km SE of Teng Chong (24°50'N,
98°47’E), collected 4-7 November 1998 by C. Griswold, D. Kavanaugh, C-L. Long, holotype and |
male and 2 female paratypes in KIZ, remainder in CAS; two male, one female paratypes, as above, but
pass over Gaolingshan at 2300 m, Luoshuidong, 28 air km E of Teng Chong (24°57'N, 98°45’E),
26-31 October 1998, C. Griswold, D. Kavanaugh, male and female in KIZ, female in CAS.

DIAGNOSIS. — Vieteuma topali (Vietnam) is much smaller than the present species and has 26
segments in both sexes; V. hubeiensis (Hubei, China) is depigmented and eyeless, whilst the present
species has both eyes and pigment.

DESCRIPTION OF MALE. — Twenty-eight segments. Total length about 15 mm, width 1.5 mm;
20-22 large, regular, well-pigmented ocelli in subtriangular eyepatch. Segments subcylindrical, seg-
mental shoulders weakly developed, setae long, acute. Color medium tan-brown, usual darker mark-
ings of purplish brown. Pregonopodal legs enlarged; coxae of legpair 7 with mesoposterior angles
drawn out into short thornlike processes, slightly excavate on posterior surfaces. Anterior gonopods
(Fig. 1A, B) with long, paired sternal processes; angiocoxites cupped, with serrate margins, two
densely fimbriated pseudoflagellar branches, additional fimbriated areas; colpocoxites narrow, api-
cally hooked; telopodites single-articled, narrow, with apical setae, inserted laterally but curving pos-
teriorly. Posterior gonopods (Fig. 1C) with small colpocoxites; telopodites with prefemur complexly
lobed on anterior side, lobes with specialized, shortened, thornlike setae; femora small, elongate, re-
curved. Legpair 10 with coxal glands, otherwise unmodified. Legpair 11 with coxal glands; trochan-
ters with sharp, mesal processes, these also on legpair 12 and gradually diminishing back to legpair 16.

DESCRIPTION OF FEMALE. — Nonesexual characters as for male. Cyphopods as in Fig. 1D, sec-
ond legpair vestigial, second sternum reduced.

ETYMOLOGY. — The name honors C-L. Long, one of the collectors.

Notes. — Vieteuma longi is syntopic with Nepalella griswoldi, n. sp., and N. magna, n. sp., both
described below. The type localities are in severely disturbed, closed-canopy forests dominated by
trees of the families Fagaceae, Lauraceae and Theaceae.

Suborder Heterochordeumatidea Shear 2000
Superfamily Heterochordeumatoidea Pocock 1894
Family Megalotylidae Golovatch 1978
Genus Nepalella Shear 1987

Nepalella now consists of 18 species (including the four new ones below) from Nepal, Thailand,
Myanmar and southern China (Shear 1987, 1999a; Mauriés 1988). As with the other megalotylid ge-
nus, Megalotyla Golovatch, the anterior gonopods are reduced to a sternocoxal plate, from which two
lateral pronglike pieces rest in lateral grooves on the anterior faces of the posterior gonopod
colpocoxites. Because of seeming similarities in the gonopod plan, megalotylids have always been
considered close to the Conotylidae, the one discordant character being the presence of coxal glands
on the eleventh legpair in the megalotylids (a plesiomorphic character). In this way megalotylids are

66 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES
Volume 53, No. 6

FiGurE 1. Vieteuma longi, n. sp. A. Anterior gonopods, anterior view. B. Anterior gonopods, posterior view. C. Posterior
gonopods, anterior view. D. Segment 2 of female, anterioventral view. Angiocoxite, a; colpocoxites, ¢; femur, f; prefemur, pf;
sternal process, s; second sternite, s2; telopodite, t.

similar to the Diplomaragnidae (like them, exclusively Asian). Diplomaragnids have a strange
gonopod structure: the telopodites of the anterior gonopods are retained (though reduced; this makes
the exact homology of these structures questionable) and articulating with a small, T-shaped sternum,
pass dorsally to the posterior gonopods and are received in grooves or partially closed pockets on the
posterior sides of the posterior gonpod coxites (Shear 1999b). Mauriés (1988) drew attention to a pre-
viously unnoticed feature of Nepalella: the anterior gonopod coxosternite has a short, platelike pro-
jection which extends dorsally between the coxae of the posterior gonopods. This is an intriguing
similarity between Nepalella and diplomaragnids, not found in any other chordeumatidan millipeds.

Diplomaragna Attems is the only genus in its family, but contains more than 30 described spe-
cies, with the rate of description of new species unabated since 1990. Nepalella likewise promises to
be a very speciose genus. What is needed to examine any possible relationship between the two genera
is material from central and northern China, where, if intermediate forms exist, they will most likely
be found.

SHEAR: CHORDEUMATIDAN MILLIPEDS FROM CHINA 67

Enough is now known about Nepalel/la species to be able to describe the general plan of both the
male and female genitalia. As already mentioned, the anterior gonopods are strongly reduced, consist-
ing only of a sternocoxal plate with a median lamellate process, and two lateral spikes. The posterior
gonopods bear large coxites that are fundamentally bipartite, especially when seen in anterior view.
Either of these divisions may be branched or simple; the lateral division is often in the form ofa broad,
flat plate turned with its axis parallel to the body midline. Posteriorly, there is at least one branch cov-
ered with fine cuticular fimbriae, and the entire posterior surface of the coxite may appear densely
hairy. The telopodites are typically reduced to a prefemur and femur, the latter turned sharply dorsad;
the whole telopodite may be quite small. In typical conotyloids and heterochordeumatoids, the femur
is swollen with gland tissue; the gland may debouche through a canal in the coxite. There is no sign of
that adaptation in Nepalella, indeed the tendency seems to be for the femur to become smaller; a simi-
lar trend is found in diplomaragnids, which also appear to lack the glands. The coxae of legpair 10 bear
large, spermatophore-forming glands, and the coxae themselves frequently (but not always) carry
well-developed processes; this appears to be usual in the Himalayan species, but the Chinese forms so
far discovered have essentially unmodified coxae 10. The telopodites of the 10th legs are of normal
proportions, but often the whole telopodite is notably smaller than that of the succeeding legs. A gland
may be present on coxa 11, or just a small vestigial homolog, sometimes reduced to a shallow depres-
sion. Prefemur | 1 usually has a dorsally directed hook. The pregonopodal legs are markedly crassate
compared to the others, and may have modifications of the prefemora and femora; this differs a great
deal from species to species.

The female genitalia are of considerable interest and of obvious taxonomic utility, contrary to the
situation in many milliped groups where gender differences between species may be subtle. Many
species have a strong clavate process on the operculum, and the valves may bear an array of teeth and
lamellae. Frequently the cuticle surrounding the valves is sclerotized and modified; two lateral cuticu-
lar processes seem to be developed either from this cuticle or from lateral extensions of the second
sternite. The distinctiveness of the female genitalia in this genus makes it acceptable to describe spe-
cies when only females are available.

Nepalella griswoldi, new species
Figure 2

MATERIAL EXAMINED. — Male holotype, 3 female paratypes from China, Yunnan Prov.,
Baoshan Prefecture, pass over Gaolingshan at 2300 m, Luoshuidong, 28 air km E of Teng Chong
(24°57'N, 98°45’E), 26-31 October 1998, C. Griswold, D. Kavanaugh, holotype and | paratype in
KIZ, 2 paratypes in CAS; male, 2 female paratypes as above, but pass over Gaolingshan at 2100 m,
Nankang, 36 air km SE of Teng Chong (24°50'N, 98°47'E), collected 4-7 November 1998 by C.
Griswold, D. Kavanaugh, C-L. Long, female paratype in KIZ, male and female in CAS.

DIAGNOSIS. — The posterior gonopod coxites of this species resemble those of NV. kavanaughi, n.
sp., but the posterior branch is densely fimbriated in N. griswoldi.

DESCRIPTION OF MALE. — Total length about 16 mm, width 2.2 mm. Thirty subcylindrical seg-
ments with well-developed dorsolateral shoulders; segmental setae short and acute. Ocelli 25, round
and well-pigmented, in triangular eyepatch. Color medium brown, mottled darker purplish brown in
usual chordeumatidan pattern. Legpairs 3—7 enlarged; femur 4 with distal knob subtending distinct
depression on mesal side. Anterior gonopods (Fig. 2A) as usual for Nepalella. Posterior gonopods
(Fig. 2A, B) with large, robust coxites apically divided, lateral branches flattened, turned with broad
planes parallel to body axis, mesal branches acute, curved, bearing indistinct internal channels; large,
thick processes dorsally directed from base of lateral branches; posterior branches and most of poste-
rior surfaces covered in fine cuticular fimbriae; telopodites of normal size but prefemora notably re-
duced. Legpair 10 with coxal glands, coxae laterally elongate, prefemora swollen, with dorsally

68 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES
Volume 53, No. 6

FiGurE 2. Nepalella griswoldi, n. sp. A. Gonopods, anterior view. B. Right posterior gonopod coxite, posterior view. C.
Cyphopods, ventral view (setae not shown on right side). Anterior gonopods, agp; colpocoxites, ¢; femur, f; prefemur, pf; re-
ceptacle process, rp.

SHEAR: CHORDEUMATIDAN MILLIPEDS FROM CHINA 69

directed proximal knobs. Legpair 11 with coxal glands replaced by convex area of finely papillate
transparent cuticle, prefemora with dorsally directed proximal hooks.

DESCRIPTION OF FEMALE. — Nonsexual characters as in male. C yphopods (Fig. 2C) with projec-
tions from valves, receptacle drawn out into blunt process.

ETYMOLOGY. — The species epithet honors Charles Griswold.

NoTEs. — The vestigial gland of legpair 11 has been noted and iilustrated for N. birmanica by
Maurieés (1988). Nepallela birmanica in several ways bridges the morphological gap between the Chi-
nese and the Himalayan species of the genus. This species is syntopic with Vieteuma longi, n. sp., de-
scribed above; see the habitat notes under the description of the preceding species.

Nepalella pianma, new species
Figure 3

MATERIAL EXAMINED. — Male holotype from China, Yunnan Prov., Nujiang Pref., native forest
at 2500 m elevation in Gaolingshan at 9.5 road km ESE of Pianma (25°59’'N, 98°40’E), collected
13-18 October 1998 by C. Griswold, D. Kavanaugh, and C-L. Long, deposited in KIZ.

DIAGNOSIS. — Distinct from all previously described species of Nepalella in the extreme reduc-
tion in size of the posterior gonopod telopodites.

DESCRIPTION OF MALE. — Total length about 14 mm, width 1.5 mm. Thirty segments with mod-
erately developed dorsolateral shoulders, segmental setae short, blunt. Eyepatch triangular, with
24-25 well-formed, pigmented ocelli. Color uniform light brown.

Pregonopodal legs crassate, but without further modifications. Anterior gonopods (Fig. 3A) as
usual for the genus; posterior gonopods (Fig. 3A, B) with large coxites bearing arcuate lateral excava-
tions lined by small teeth, posteriorly with single moderately developed fimbriate branch, telopodites
much reduced. Legpair 10 with large glands but coxae not conspicuously enlarged or modified;
legpair 11 with very small, probably vestigial, glands, coxae not enlarged, prefemora with dorsally di-
rected posterioproximal hooks.

Female unknown.

ETYMOLOGY. — The species epithet is a noun in apposition, after the type locality.

FiGuRE 3. Nepalella pinama, n. sp. A. Gonopods, anterior view. B. Right posterior gonopod coxite, posterior view. Anterior
gonopods, agp; colpocoxites, ¢; femur, f; prefemur, pf.

70 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES
Volume 53, No. 6

NOTES. — This species is syntopic with Nepallela kavanaughi, n. sp., described below. The
Pianma forest is similar to that already described, with oaks, tanbarks, chinquapins, etc., predominat-
ing, but at this higher-elevation site maples, ashes and giant rhododendrons were more prominent.
This forest was only slightly disturbed by human activity.

Nepalella kavanaughi, new species
Figure 4

MATERIAL EXAMINED. — Male holotype, male and four female paratypes from China, Yunnan
Prov., Nujiang Pref., native forest at 2500 m elevation in Gaolingshan at 9.5 road km ESE of Pianma
(25°59'N, 98°40’E), collected 13—18 October 1998 by C. Griswold, D. Kavanaugh, and C-L. Long,
holotype and 2 female paratypes in KIZ, male and 2 female paratypes in CAS.

DIAGNOSIS. — The posterior gonopod coxites resemble those of N. griswoldi, but are markedly
less fimbriate posteriorly; the dorsally enlarged prefemora of legpairs 3—7 are unique in the genus, as
is the lack of a prefemoral process on legpair 11.

DESCRIPTION OF MALE. — Total length about 14 mm, width |.8—2.0 mm. Thirty subcylindrical
segments with strongly developed dorsolateral shoulders, segmental setae moderately long, acute.
Color even medium brown with darker markings on anterior segments and head. Ocelli 25, black, in
subtriangular eyepatch. Legpairs 3—7 crassate, prefemora with swollen dorsal sides (Fig. 4A). Ante-
rior gonopods typical for genus (Fig. 4B). Posterior gonopod coxites (Fig. 4B, C) with two anterior
branches; single posterior branch blunt, not fimbriated; telopodites of normal size for genus. Legpair
10 (Fig. 4D) with coxal glands, coxae slightly elongate above gland, telopodites of reduced size but
complete and proportionate. Legpair 11 (Fig. 4D) with vestigial glands as described for N. griswoldi,
but lacking prefemoral processes.

DESCRIPTION OF FEMALE. — Nonsexual characters as in male. Cyphopods (Fig. 4E) with valvu-
lar processes reduced, receptacle process unusually long, well-sclerotized; entire cyphopod complex
encased in sclerotized extensions of sternum 2, extensions drawn out ventrally into blunt blades.

NOTES. — This species is syntopic with N. pianma, n. sp., described above. See the habitat notes
under that species.

Nepalella magna, new species
Figure 5

MATERIAL EXAMINED. — Female holotype, 3 female paratypes from China, Yunnan Prov.,
Baoshan Prefecture, pass over Gaolingshan at 2300 m, Luoshuidong, 28 air km E of Teng Chong
(24°57'N, 98°45'E), 26-31 October 1998, C. Griswold, D. Kavanaugh, holotype and | paratype in
KIZ, 2 paratypes in CAS.

DIAGNOSIS. — Distinct from other Chinese Nepale/ia in its large size (about 30% larger than any
known species) and in details of the cyphopods, as depicted in Figure 5.

DESCRIPTION OF FEMALE. — Total length about 33 mm, width 3.0 mm. Thirty subcylindrical
segments with moderately developed shoulders; setae short, acute. Twenty-five large, black ocelli
form triangular eyepatch. Color dark brown, heavily marked in typical chordeumatidan fashion with
darker, purplish brown.

Cyphopods (Fig. 5) with distinct valvular processes; receptacle processes expanded distally,
curving posteriorly over valves. Sternum 2 with large, crenulate knob extending between pairs of
valves, laterally and posteriorly around cyphopods, bearing long, thin, acute processes. In undissected
animals, these processes lie between third leg coxae and third sternite.

Male unknown.

ETYMOLOGY. — The species epithet is an adjective referring to the great size of this species.

SHEAR: CHORDEUMATIDAN MILLIPEDS FROM CHINA 71

FiGure 4. Nepalella kavanaughi, n. sp. A. Legpair 3 of male, posterior view. B. Gonopods, anterior view. C. Right posterior
gonopod coxite, posterior view. D. Legpairs 10 (foreground) and 11 of male, anterior view. E. Cyphopods, ventral view. Ante-
rior gonopods, agp; colpocoxites, ¢; femur, f; prefemur, pf; receptacle process, rp; numbers indicate legpair numbers.

72 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES
Volume 53, No. 6

FIGURE 5. Cyphopods of Nepalella magna, n. sp., ventral view. Receptacle process, rp; sternal process, sp.

NOoTEs. — I have taken the unusual step of describing this species without males because of the
distinctiveness of the cyphopods, which bear the characteristic receptacle process found in Nepalella,
and the unusually large size, which will make it possible to recognize males when they are collected.
This species is syntopic with Nepalella griswoldi at their common type locality, and with Vieteuma
longi. The habitat was described under the latter species.

LITERATURE CITED

GOLOVATCH, S. I. 1984. Contributions to the milliped fauna of Vietnam (Diplopoda) II. Acta Zoologica
Hungarica 30:53-77.

MAURIES, J.-P. 1988. Myriapodes du Népal. II. Diplopodes craspedosomides nouveaux de |’ Himalaya et de la
région indo-malaise (Craspdesomidea et Chordeumidea). Revue Suisse Zoologie 95:3—-49.

MAURIES, J.-P. AND M. NGUYEN DUY-JAQUEMIN. 1997. Nouveaux craspedosomides et glyphiulides cavernicoles
de Chine (Diplopoda). Mémoires Biospéleologique 24:49—62.

SHEAR, W. A. 1987. Chordeumatid Diplopoda from the Nepal Himalayas, II. Courier Forschungs-Institut
Senckenberg 93:229-240.

. 1999a. A new troglobitic milliped of the genus Nepalella from China (Diplopoda, Chordeumatida,

Megalotylidae). Myriapodologica 6:1—10.

. 1999b. The milliped genus Diplomaragna confirmed for Taiwan, with the description of a new species
(Diplopoda, Chordeumatida, Diplomaragnidae). Myriapodologica 6:1 1—18.

WANG, D. AND J.-P. MAURIES. 1996. Review and perspective of study on myriapodology of China. Mémoires
Museum National d’Histoire Naturelle (Paris) 169:81—99.

PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES

Volume 53, No. 7, pp. 73-86, 5 figs., 2 tables. July 1, 2002

by
Aaron M. Bauer 20 02
Department of Biology, Villanova University,

800 Lancaster Avenue, Villanova, Pennsylvania 19085 \ij3¢)- ,, re
; ; ae ‘ NOY HANS Ue 3 ig)
email: aaron.bauer@villanova.edu

a SESE

Two new species of the gekkonid lizard genus Cyrtodactylus are described on the basis of
material collected in Popa Mountain Park and Alaungdaw Kathapa National Park in
north central Myanmar. The species from Popa Mountain is highly distinctive and resem-
bles members of the Indian-Sri Lankan subgenus Geckoella in external appearance. It is
characterized by its short digits, large number of midventral scales, series of 8 precloacal
pores and absence of precloacal groove, femoral pores and enlarged femoral scales. The
larger species, from Alaungdaw Kathapa, is diagnosed by its long digits and tail, absence of
precloacal groove, series of 9-11 precloacal pores separated from a series of 11-12 femoral
pores on each side, enlarged subcaudal plates, and pattern of 6-8 pairs of more-or-less
well-defined, dark dorsal blotches between the nape and sacrum. The larger of the two spe-
cies may be allied to C. consobrinus and other large-bodied southeast Asian members of the
genus. The gekkonid fauna of Myanmar is more diverse than previously recognized and in-
cludes as many as 12 species of Cyrtodactylus.

Both the tropical and temperate parts of Eurasia are occupied by gekkonid lizards that lack ex-
panded toe pads and are characterized by a “bent-toed” appearance. The taxonomic history of these
geckos is complex. Prior to the work of Underwood (1954) these taxa were typically lumped, along
with other padless geckos, into the genus Gymnodactylus. Underwood (1954) restricted this name to
South American forms, resurrecting Cyrtodactylus Gray for Old World taxa. Palearctic bent-toed
geckos have subsequently been revised by Szezerbak and Golubev (1977, 1984, 1986), who demon-
strated that fundamental differences exist between these taxa and tropical Asian Cyrtodactylus. The
nomenclature of this group has been problematic (Kluge 1985; Mees 1987) and robust phylogenetic
hypotheses at both the inter- and intrageneric levels are lacking. Kluge (1991, 1993, 2001) recognized
two genera of tropical Asian bent-toed geckos: Geckoella, occurring in Peninsular India and Sri
Lanka, and Cyrtodactylus, distributed from south Asia to the western Pacific, as far as the Solomon Is-
lands. Generic composition in Cyrtodactylus has remained unstable, chiefly with respect to certain
taxa occurring in northern India and adjacent regions (e.g., Kluge 1993, 2001; Bauer and Henle 1994;
Khan and Résler 1999; Résler 2000; Wells 2002). R6sler (2000) placed all Palearctic taxa of ques-
tionable affinities (including Gonydactyluys martinstollii Darevsky et al., 1997, G. markuscombaii
Darevsky et al., 1997, and G. nepalensis Schleich and KAstle, 1998) in the genus Cyrtopodion, but re-
garded Geckoella as a subgenus of Cyrtodactylus. Under this interpretation 63 species of
Cyrtodactylus are currently recognized [Résler 2000, 2001; Das and Lim 2000; Wells 2002;
Cyrtodactylus mansarulus Duda and Sahi, 1978, although treated by Rosler (2000) as a
Cyrtodactylus, is probably correctly assigned to Cyrtopodion sensu lato]. Numerous additional un-

74 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES
Volume 53, No. 7

described taxa are known to exist (Bauer and Henle 1994; R. Giinther, pers. comm.), making
Cyrtodactylus one of the most speciose of all gecko genera.

Smith (1935) recorded six species of Cyrtodactylus from Burma: C. feae, C. consobrinoides, C.
khasiensis, C. variegatus, C. oldhami, and C. peguensis. Hundley (1964) listed seven species, omit-
ting C. feae but including one species referable to Cnemaspis. He also added C. rubidus, an Andaman
Islands endemic to the fauna, presumably on the basis of material from the Cocos Group, the only part
of the Andamans under the administration of Myanmar. Das (1997) and Das and Lim (2000) also indi-
cated the occurrence of C. pulchellus from the country. I here describe two hitherto unknown mem-
bers of the genus collected by the Myanmar Herpetofaunal Survey in the Sagaing and Mandalay
Divisions of north central Myanmar.

MATERIALS AND METHODS

The following measurements were taken with Brown and Sharpe Digit-cal Plus digital calipers
(to the nearest 0.1 mm): snout-vent length (SVL; from tip of snout to vent), trunk length (TrunkL; dis-
tance from axilla to groin measured from posterior edge of forelimb insertion to anterior edge of
hindlimb insertion), crus length (CrusL; from base of heel to knee); tail length (TailL; from vent to tip
of tail), tail width (TailW; measured at widest point of tail); head length (HeadL; distance between
retroarticular process of jaw and snout-tip), head width (HeadW; maximum width of head), head
height (HeadH; maximum height of head, from occiput to underside of jaws), ear length (EarL; lon-
gest dimension of ear); forearm length (ForeaL; from base of palm to elbow); orbital diameter (OrbD;
greatest diameter of orbit), nares to eye distance (NarEye; distance between anteriormost point of eye
and nostril), snout to eye distance (SnEye; distance between anteriormost point of eye and tip of
snout), eye to ear distance (EyeEar; distance from anterior edge of ear opening to posterior corner of
eye), internarial distance (Internar; distance between nares), and interorbital distance (Interorb; short-
est distance between left and right supraciliary scale rows).

Scale counts and external observations of morphology were made using a Nikon SMZ-10 dis-
secting microscope. Radiographic observations were made using a Faxitron closed cabinet X-ray sys-
tem. Comparisons were made with museum material in the collections of the California Academy of
Sciences (CAS), United States National Museum (USNM), and the Museum of Comparative Zoology
(MCZ), as well as original published descriptions and descriptions provided in broader faunal and tax-
onomic treatments (e.g., Smith 1935; Szczerbak and Golubev 1986; Hikida 1990; Ulber 1993;
Darevsky and Szezerbak 1997; Das 1997; Das and Lim 2000).

Geographic coordinates were recorded by the original collectors from a Garmin 12 GPS (datum
WGS 84).

SYSTEMATICS

REPTILIA: SQUAMATA: GEKKONIDAE

Cyrtodactylus brevidactylus, sp. nov.
Figs. 1,2

HOLOTYPE. — CAS 214104 (Field number JBS 4210), adult male; Popa Mountain Park, Kyauk
Pan Tawn Township, Mandalay Division, Myanmar (20°57'56.8”N, 95°14'18.8”E); collected by
Htun Win, Thin Thin, Sai Wunna Kyi and Hla Tun, 26 March 2000.

PARATYPES. — CAS 214105 (Field number JBS 4211), adult female; data as for holotype. CAS
214120 (Field number JBS 4231), adult female; Popa Mountain Park, Kyauk Pan Tawn Township,

BAUER: NEW SPECIES OF CYRTODACTYLUS (5

Mandalay Division, Myanmar (20°59'00.2"N,
95°16'12.8"E); collected by Htun Win, Thin Thin,
San Lwin Oo, Sai Wunna Kyi and Hla Tun, 27
March 2000.

ETYMOLOGY. — From the Latin brevis, short,
and the latinized Greek dactylus, finger, in recogni-
tion of the very short digits of this species. The epi-
thet is an adjective in the nominative singular.

DEFINITION. — A moderately sized Cyrto-
dactylus, snout-vent length 71-88 mm; body robust,
limbs, and especially digits, short; two pairs of en-
larged postmental scales, members of each pair in
broad contact with one another; dorsal scales con-
sisting of 27 rows of enlarged keeled tubercles, each
surrounded by smaller, conical scales, yielding a
very rugose appearance; 45 ventral scales between
lowest rows of enlarged flank tubercles; no
precloacal groove, 8 precloacal pores in a single se-
ries in both males and females, no femoral pores or
enlarged femoral scales. Eleven subdigital lamellae
beneath 4th toe of pes distal to digital inflection.
Subcaudal scalation (based on original portion of
holotype tail) uniform and granular, without en-
larged midventral plates. Dorsal pattern of 3-4 large
dark dorsal blotches between nape and sacrum. Head
chocolate brown with white markings, including a
Y-shaped mark on the nasal, frontal and parietal sur-
faces.

DESCRIPTION (based on _ holotype, CAS
214104). — Adult male with midventral incision
from tissue removal. Snout-vent length 71.4 mm. FicuRE 1. Holotype of Cyrtodactylus brevidactylus,
Head short (HeadL/SVL ratio 0.20), relatively wide Ne ay siahrai on Boe ea fa
(HeadW/Headl. ratio 0.64), not depressed (HeadH/ Manna. Notte shor, set regsnaatd il hot
HL ratio 0.39), distinct from neck. Lores and
interorbital region inflated, canthus rostralis promi-
nent. Snout short (SnEye/HeadL ratio 0.34); longer than eye diameter (OrbD/SnEye ratio 0.61);
scales on snout and forehead rounded, granular, becoming heterogeneous posteriorly; scales on snout
smaller than those on occipital region. Eye relatively small (OrbD/HeadL ratio 0.20); pupil vertical
with crenelated margins; supraciliaries short, conical. Ear opening oval, relatively large (EarL/HeadL
ratio 0.05); greatest diameter vertical; eye to ear distance greater than diameter of eyes (EyeEar/OrbD
ratio 1.25). Rostral approximately 2/3 deep (2.0 mm) as wide (3.0 mm), incompletely divided dor-
sally by weakly developed rostral groove; two enlarged supranasals separated by a single, roughly
pentagonal internasal; rostral in contact with supralabial I, supranasals, and internasal; nostrils oval,
each surrounded by supranasal, rostral, first supralabial, enlarged postnasal and narrow crescentic na-
sal; nasal in turn bordered by three small postnasal scales; two rows of scales separate orbit from
supralabials. Mental subtriangular, much wider (3.2 mm) than deep (1.8 mm); two pairs of enlarged
postmentals, anteriormost approximately 3—4 times larger than posterior, posterior approximately
3—5 times larger than adjacent chinshields; each anterior postmental bordered anteriorly by mental,
medially by other anterior postmental, laterally by first infralabial, and posteriorly by an enlarged lat-

76 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES
Volume 53, No. 7

FIGURE 2. Living specimen of Cyrtodactylus brevidactylus, sp. nov. (CAS 214120, paratype) from Popa Mountain Park,
Myanmar. Photo by Hla Tun.

eral chinshield, a small granule, and posterior postmental; posterior postmentals each bordered later-
ally and posteriorly by series of 5-6 granules. Infralabials bordered by a row of scales slightly larger
than more posterior throat granules. Supralabials (to midorbital position) 8 (right)-10 (left);
supralabials to angle of jaws 11 (right)—12 (left); infralabials 11: interorbital scale rows across nar-
rowest point of frontal 17.

Body stout, elongate (TrunkL/SVL ratio 0.40) with very weakly developed, non-denticulate
ventrolateral folds. Dorsal scales heterogeneous, conical: regularly arranged large tubercles extend-
ing from posterior border of orbits and temporal region on to back; each tubercle bearing ridges, the
most prominent forming a keel on the anterior-facing surface; enlarged tubercles surrounded by ro-
settes of smaller conical scales of varying sizes, the largest being anterior to the tubercle; tubercles be-
coming smaller and less prominently ridged on flanks: largest tubercles in approximately 27 rows at
midbody. Ventral scales smaller than dorsal, weakly conical, posteriorly oriented and subimbricate;
somewhat larger on chest and in precloacal region than elsewhere: midbody scale rows across belly to
lowest row of enlarged lateral tubercles 45; gular region with strongly heterogeneous scalation; larger
conical scales among smaller, mostly rounded, granules. Precloacal pores in a single series of 8 witha
narrow poreless scale separating two continuous rows of 4 on each side; lateralmost scales flanked by
an enlarged, slightly dimpled scale; scale row immediately posterior to pores much enlarged, but
without evidence of glands or pores. No femoral pores or enlarged femoral scales; no precloacal
groove. Scales on palm and sole smooth, rounded: scales on dorsal aspects of limbs keeled, similar to
dorsal scales, with larger, conical tubercles interspersed among smaller scales.

Fore and hindlimbs relatively short, stout; forearm short (ForeaL/SVL ratio 0.16); tibia short
(CrusL/SVL ratio 0.18); digits very short, strongly inflected at basal interphalangeal joints, all bearing
robust, slightly recurved claws; subdigital lamellae narrow, rounded, enlarging somewhat distally,
without scansorial surfaces; scansors distal to raised circular scale at digital inflection and not includ-
ing ventral claw sheath: 8-7-10-9-8 (manus), 6-9-11-11-10 (pes); median scales proximal to digital in-
flection not or only slightly larger than adjacent granules; interdi gital webbing absent. Relative length
of digits (manus; measurements in mm in parentheses): IV (4.78) > III (4.47) > II (4.24) > V (3.01) >1
(2.80); (pes): IV (5.77) > V (5.11) > III (4.49) > II (3.74) > I (2.82).

Mostly regenerated tail short, very stout, constricted at base and with an abruptly narrowing tip;
regenerated tail much shorter than snout-vent length (TailL/SVL ratio 0.48): original portion of tail

BAUER: NEW SPECIES OF CYRTODACTYLUS qi

with transverse rows of enlarged, conical tubercles, regularly spaced with approximately 6 rows of
smaller scales intervening; ventral scales more uniform in size, keeled and conical, but more rounded
than dorsals. Regenerated portion of tail with more-or-less uniform, rounded, rugose scales; without
tubercles; 4 (right) or 5 (left) enlarged, striated, conical postcloacal spurs on each side of tailbase.

Osteology. Frontal bone extremely wide, with very prominent supraorbital ridges. Parietal bones
paired. Premaxillary teeth loci 11, approximately 40 teeth on each maxillary bone, 47 on each dentary.
Phalangeal formulae 2-3-4-5-3 for manus and 2-3-4-5-4 for pes; basal phalanges of manus imbedded
in tissue of palm. Presacral vertebrae 26, including 3 anterior cervical (without ribs), | lumbar, and 2
sacral vertebrae; 5 pygal and 3.5 post pygal caudal vertebrae to point of regeneration (0.5 post-pygal
vertebrae in both paratypes). Two pairs of cloacal bones present, medial pair crescentic, lateral pair
nodular, lying beneath the raised cloacal spurs (cloacal bones lacking in female paratypes).
Endolymphatic sacs not enlarged extracranially.

Coloration (in preservative). Base color a pale grayish brown to white. Boldly marked with nu-
merous chocolate brown markings, each outlined sequentially by a relatively thick, dark brown bor-
der and a much thinner, whitish border, the latter only about one scale width in thickness. Dorsum of
head predominantly chocolate brown with a paler Y-shaped marking; base of Y extending posteriorly
from a pale, irregular marking on intercanthal region, bifurcating behind orbits and angling laterally
towards upper margin of ear to fuse with whitish region extending from the jaw to the nape and
forelimb insertion. Another light line extends from supraciliary scales backward to join the transverse
line above the ear. A diffuse, pale marking present at anteroventral margin of orbit. Labial scales white
with chocolate brown markings.

Dorsum marked by a series of 3-4 large, roughly symmetrical blotches between the nape and sa-
crum. A similar marking, but consisting of separate left and right blotches, on tail base. Lateral surface
with a series of spots, including three behind the ear, one over the scapular region, and one or more be-
tween the lateral borders of each of the enlarged dorsal blotches. Markings along the ventrolateral
margin of the body narrow and sometimes barlike. Limbs marked dorsally by a series of dark trans-
verse bars that extend onto the edges of the ventral surfaces. Venter cream white tinged by the light
brown speckling of individual scales. Tail mottled gray-brown with darker irregular transverse mark-
ings.

Color in life essentially identical (Fig. 2).

VARIATION. — Comparative mensural data for the holotype and paratypes is presented in Table
1. Both paratypes similar to holotype in most respects except as noted. CAS 214120: Adult female
with midventral incision from tissue removal. Three small internasal scales present; supralabial scales
to midpoint of orbit 8, to angle of jaw 10 (right)—11 (left); infralabials 11. Distal portion of digit II, left
manus missing. Number of lamellae distal to enlarged scale at metaphalangeal inflection 6-7-9-9-6
(manus), 8-9-1 1-10-11 (pes). Precloacal pores as in holotype; cloacal spurs 3-4, much smaller than in
holotype. Regenerated tail rounded without tapered tip, almost uniform dark brown. Dorsal pattern
similar to holotype but base color a darker pinkish brown and markings darker and bolder; dark mark-
ings on either side of tail base fused at midline to form a complete band. Light lines on head in this and
second paratype partially disrupted in temporal region. Venter beige, more heavily pigmented than
holotype; throat with scattered dark blotches. CAS 214105: Adult female. One internasal scale;
supralabial scales to midpoint of orbit 8, to angle of jaw 10 (right)—11 (left); infralabials 10
(left)—1 1(right). Number of lamellae distal to enlarged scale at interphalangeal inflection 7-7-10-9-8
(manus), 8-9-1 1-10-10 (pes). Precloacal pores as in holotype; cloacal spurs 3-4, much smaller than in
holotype. Light markings on head less well defined than in holotype, not continuous. An extra band
anterior to sacral blotch formed from fusion of smaller dorsolateral spots. Postcloacal band and tail as
in CAS 214120. Limbs more darkly pigmented than in holotype. Ventral coloration beige with scat-
tered brown marks, forming faint reticulations on margins of throat.

78 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES
Volume 53, No. 7

TABLE 1. Mensural data for the type series of Cyrtodactylus brevidactylus, sp. nov.
Abbreviations as in Materials and Methods, all measurements in mm.

CAS 214104 CAS 214120 CAS 214105

holotype paratype paratype
Sex male female female
SVL (ales! 88.0 84.3
ForeaL LS) 12.9 13.3
CrusL [Sel 14.4 16.3
TailL (entire) 35.0 23.6 19.3
TailL (portion regenerated) 24.0 17.0 14.6
TailW 9.0 10.9 11.3
TrunkL 28.6 39.8 39.5
HeadL 20.2 22.8 24.0
Head W 12.9 S35 14.9
HeadH 7.8 9.5 10.5
OrbD 4.] S22 57)
EyeEar ae 5.6 59
SnEye 6.8 8.0 S.2
NarEye 4.6 5.4 4.9
Interorb ES 9.0 9.7
BarL el 1.3 3)
Internar 1.8 2.6 1.9

COMPARISONS. — Cyrtodactylus brevidactylus may be distinguished from all congeners on the
basis of its stout body, short digits, series of 8 precloacal pores and no precloacal groove, femoral
pores or enlarged femoral scales, highly tuberculate dorsal scalation, and dorsal color pattern, includ-
ing Y-shaped marking on dorsum of head. The condition of precloacal and femoral pores 1n males has
traditionally been widely used to distinguish members of the genus Cyrtodactylus (e.g., Darevsky and
Szezerbak 1997). On this basis C. brevidactylus may be distinguished from the following species by
the absence of a precloacal groove: C. annulatus (Taylor, 1915), C. cavernicolus (Inger and King,
1961), C. marmoratus (Gray, 1831), C. papuensis (Brongersma, 1934), C. philippinicus
(Steindachner, 1867), C. pubisulcus Inger, 1958, C. pulchellus Gray, 1827, C. rubidus (Blyth, 1860),
C. sadleiri Wells and Wellington, 1984; from the following species by the absence of femoral pores
and/or enlarged femoral scales: C. abrae Wells, 2002, C. agusanensis (Taylor, 1915), C. angularis
(Smith, 1921), C. baluensis (Mocquard, 1890), C. biordinis Brown and McCoy, 1980, C.
brevipalmatus (Smith, 1923), C. condorensis (Smith, 1920), C. consobrinoides (Annandale, 1905),
C. consobrinus (Peters, 1871), C. darmandvillei (Weber, 1890), C. derongo Brown and Parker, 1973,
C. feae (Boulenger, 1893), C. fraenatus (Giinther, 1864), C. fumosus (Miller, 1895), C.
gubernatorius (Annandale, 1913), C. interdigitalis Ulber, 1993, C. intermedius Smith, 1917, C.
irregularis (Smith, 1921), C. jarujini Ulber, 1993, C. lateralis (Werner, 1896), C. loriae (Boulenger,
1898), C. louisiadensis (de Vis, 1892), C. malcolmsmithi (Constable, 1949), C. mimikanus
(Boulenger, 1914), C. novaeguineae (Schlegel, 1844), C. oldhami (Theobald, 1876), C. papilionoides
Ulber, 1993, C. peguensis (Boulenger, 1893), C. quadrivirgatus Taylor, 1962, C. redimiculus King,
1962; C. sworderi (Smith, 1925), C. tiomanensis Das and Lim, 2000, C. tuberculatus (Lucas and
Frost, 1900), C. variegatus (Blyth, 1859), C. wetariensis (Dunn, 1927), and C. n. sp. Bauer, 2002 (this
paper); from the following species by the presence of precloacal pores: C. je/lesmae (Boulenger,
1897), C. laevigatus Darevsky, 1964, C. paradoxus (Darevsky and Szczerbak, 1997), C.
sermowaiensis (de Rooij, 1915), and most members of the subgenus Geckoella [C. collegalensis
(Beddome, 1870), C. deccanensis (Giinther, 1864), C. jevporensis (Beddome, 1877), C. nebulosus
(Beddome, 1870), and C. yakhuna (Deraniyagala, 1945)]; and from C. (G.) triedrus (Giinther, 1864)

BAUER: NEW SPECIES OF CYRTODACTYLUS 79

by having 8 vs. 34 precloacal pores. In compar-
ison with remaining congeners, the presence of
two (vs. one) pairs of enlarged postmental
scales distinguish the new species from C.
adleri Das, 1997, C. irianjayaensis Résler,
2001, and C. malayanus (de Roo, 1915) and
the broad contact of second postmentals (vs.
wide separation by anterior postmentals) distin-
guish it from C. e/ok Dring, 1979, C. ingeri
Hikida, 1990, C. khasiensis (Jerdon, 1870), C.
matsuii Hikida, 1990, and C. yoshii Hikida,
1990.

Furthermore, in comparison to other spe-
cies known from Myanmar, the approximately
45 midventral scale rows of C. brevidactylus is
greater than the comparable scale counts re-
corded for all other species (data from Smith,
1935 and Das, 1997): C. consobrinoides (35), C.
feae (35), C. khasiensis (30-40), C. oldhami
(34-38), C. peguensis (34-38 fide Smith 1935;
32 fide Das 1997), C. pulchellus (35), C.
variegatus (22), C. n. sp. (this paper) (27-32).
Each of these species also has a somewhat de- =
pressed habitus and differs markedly in color FiGuRE 3. Map of Myanmar illustrating the distribution of C.
pattern from the new species (longitudinal brevidactylus and & slowinskit in the central dry zone of the

: ; : 5 country. This region is bounded to the west by the Rakhine
stripes in C. oldhami, cross bands in C. Yoma (Arakan Yoma) and to the east by the montane systems
consobrinoides, C. pulchellus; more numerous _ of Indochina. Map prepared by Michelle S. Koo, California
dark markings in the remaining species). Academy of Sciences.

DISTRIBUTION. — Cyrtodactylus brevi-
dactylus is known only from Popa Mountain in
the Mandalay Division of Myanmar (Fig. 3). The relative isolation of this physiographic feature sug-
gests that the species may be a highly localized endemic, although further collecting in the central dry
zone of Myanmar will be required to confirm this.

REPTILIA: SQUAMATA: GEKKONIDAE

Cyrtodactylus slowinskii, sp. nov.
Figs. 4, 5

HOLOTYPE. — CAS 210214 (Field number JBS 2765), adult male; Sunthaik Chaung (tributary to
Hkaungdin Chaung), Alaungdaw Kathapa National Park, Monywa District, Sagaing Division,
Myanmar (22°18'48.8"N, 94°24'29.0"E); collected by J. B. Slowinski, K. D. Wiseman, J. M. Lovette
and J. V. Vindum, 11 July 1999.

PARATYPES. — CAS 210212—13, 210215—19 (Field numbers JBS 2739-40, 2766-70); data as
for holotype. CAS 215474 (Field number JBS 6517); Thabake Say Camp, Alaungdaw Kathapa Na-
tional Park, Monywa District, Sagaing Division, Myanmar (22°19'06.6"N, 94°28'32.0"E); collected
by Htun Win, Thin Thin, San Lwin Oo, and Sai Wunna Kyi, 26 May 2000. CAS 215593 (Field num-
ber JBS 6762); Thabake Say Camp, Alaungdaw Kathapa National Park, Monywa District, Sagaing
Division, Myanmar (22°18'29.4”N, 94°30'10.7”E); collected by Htun Win, Thin Thin, and San Lwin

80 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES
Volume 53, No. 7

Oo, 13 June 2000. CAS 215661 (Field number JBS
6965); Thabake Say Camp, Alaungdaw Kathapa Na-
tional Park, Monywa District, Sagaing Division,
Myanmar (22°18'06.6"N, 94°28'32.0"E); collected by
Htun Win, Thin Thin, San Lwin Oo, and Hla Tun, 29 June
2000. USNM 548141 (Field number JBS 6513), USNM
548142 (Field number JBS 6517); Swun Theik Camp,
base of pine mountain, Alaungdaw Kathapa National
Park, Monywa District, Sagaing Division, Myanmar
(22°18'50.1"N, 94°24'25.7"E); collected by Htun Win,
Thin Thin, San Lwin Oo, Sai Wunna Kyi, and Hla Tun,
30 May 2000. USNM 548143 (Field number JBS 6764);
Thabake Say Camp, Alaungdaw Kathapa National Park,
Monywa_ District, Sagaing Division, Myanmar
(22°18'29.4"N, 94°30'10.7"E); collected by Htun Win,
Thin Thin, and San Lwin Oo, 13 June 2000.

ADDITIONAL MATERIAL (27 specimens). — All
specimens from Alaungdaw Kathapa National Park,
Monywa District, Sagaing Division, Myanmar. CAS
210110—11; Trail between Thabake Say Camp and
Alaungdaw Kathapa Pagoda Q2- IS TOON:
94°28'30.7"E); collected by J. B. Slowinski, K. D. Wise-
man, J. M. Lovette, and J. V. Vindum, 6 July 1999. CAS
210205—11; data as for holotype. CAS 210220; locality
and collectors for holotype, 11 July 1999. CAS 210250;
Main road between Thabake Say Camp and Payawa
Sakhan (Elephant Camp) (22°19'09.0"N, 94°28'45.6"E):
collected by K. D. Wiseman, J. M. Lovette, and J. V.
Vindum, 13 July 1999. CAS 210265; Left bank of
Pwedon Chaung across from Payawa Sakhan (Elephant
Camp) (22°19'20.4"N, 94°29'11.3"E); collected by J. B.

FiGuRE 4. Holotype of Cyrtodactylus slowinskii, Slowinski, K.D. Wiseman, J. M. Lovette, and J. V.
National Parke Myanmar, Note the relatively smal Windum, 13 July 1999. CAS 210292; Paya Chaung
dorsal tubercles, elongate digits, and pattern of (22°18'54.7”"N, 94°28'35.5”E); collected by K. D. Wise-
paired dorsal markings. man, J. M. Lovette, and J. V. Vindum, 15 July 1999. CAS

210293—95; Paya Chaung (22° 183529)

94°28'42.8"E); collected by K.D. Wiseman, J. M.
Lovette, and J. V. Vindum, 15 July 1999. CAS 215483—92; Swun Theik Chaung, base of pine moun-
tain (22°18'50.1"N, 94°24'25.7”E); collected by Htun Win, Thin Thin, San Lwin Oo, Sai Wunna Kyi,
and Hla Tun, 30 May 2000. CAS 215711; Gon Nyin Bin (22°15'33.2"N, 94°36'09.6”E); collected by
Htun Win, Thin Thin, San Lwin Oo, and Hla Tun, 4 July 2000.

ETYMOLOGY. — The specific epithet is a patronym honoring my late friend and colleague Joseph
B. Slowinski, preeminent contributor to the herpetology of Myanmar. The epithet is a masculine noun
in the genitive case.

DEFINITION. — A large sized Cyrtodactylus, snout-vent length to 108 mm; body relatively slen-
der, digits long; one pair of enlarged postmental scales, in broad contact with one another; 19-22 rows
of flattened, weakly-keeled tubercles; 27—32 ventral scales between weakly-developed ventrolateral
folds; no precloacal groove, 9 precloacal pores (in holotype) ina single series in males only, 11 femo-

BAUER: NEW SPECIES OF CYRTODACTYLUS 81

ral pores (in holotype) on each thigh, separated by a diastema from precloacal pores. Six widened
subdigital lamellae beneath basal phalanx of 4th toe of pes, 13 narrow lamellae beneath more distal
phalanges of same toes. Subcaudal scalation with single series of enlarged midventral plates. Dorsal
pattern of 6-8 paired, dark, dorsal blotches between nape and sacrum; tail banded brown and white.

DESCRIPTION (based on holotype, CAS 210214). — Adult male, SVL 98.8 mm. Head moder-
ately long (HeadL/SVL ratio 0.27), wide (HeadW/HeadL ratio 0.72), not depressed (HeadH/HeadL
ratio 0.44), distinct from neck. Lores and interorbital region somewhat inflated, canthus rostralis not
well developed. Snout short (SnEye/HeadL ratio 0.40); longer than eye diameter (OrbD/SnEye ratio
0.80); scales on snout and forehead rounded, granular, intermixed with small tubercles posteriorly;
scales on snout much larger than those on occipital region. Eye relatively large (OrbD/HeadL ratio
0.32); pupil vertical with crenelated margins; supraciliaries short, blunt except for a few pointed
scales at posterior border of orbit. Ear opening rounded, relatively large (EarL/HeadL ratio 0.08); eye
to ear distance slightly greater than diameter of eyes (EyeEar/OrbD ratio 1.07). Rostral approximately
60% as deep (2.5 mm) as wide (4.2 mm), incompletely divided dorsally by weakly developed rostral
groove; two enlarged supranasals separated by a single, smaller, roughly hexagonal internasal; rostral
in contact with supralabial I, supranasals, and internasal; nostrils oval, laterally oriented, each in broad
contact with rostral and also surrounded by supranasal, first supralabial, and a narrow crescentic nasal
rim bordered by four postnasals; pigmented narial flap occludes posterior 2/3 of nostril; 4-5 rows of
scales separate orbit from supralabials. Mental triangular, wider (4.5 mm) than deep (3.2 mm); one
pair of enlarged postmentals, each approximately 40% size of mental; left and right postmentals in
broad medial contact with no intervening granules, bordered laterally by first infralabial and an en-
larged lateral chinshield, and posteriorly by 4—5 granular chin scales. Infralabials bordered by 1-3
rows of enlarged scales, largest anteriorly. Throat scales small, rounded, granular. Supralabials (to
midorbital position) 8; enlarged supralabials to angle of jaws 10 (left)-11 (right); infralabials 9;
interorbital scale rows across narrowest point of frontal bone 22.

Body moderately slender (TrunkL/SVL ratio 0.45) with very weakly developed, non-denticulate
ventrolateral folds. Dorsal scales heterogeneous, mostly flattened granules with regularly arranged
small, flattened, feebly-keeled tubercles extending from occipital region on to back and base of tail;
tubercles most prominent on dorsolateral surfaces, somewhat smaller and flatter on nape, along
middorsal line, and on flanks; tubercles in approximately 20 irregular longitudinal rows at midbody;
52 tubercles in paravertebral row from occiput to midsacrum . Ventral scales much larger than dorsal,
cycloid, imbricate to subimbricate; largest under thighs and between precloacal pores and vent, some-
what larger on abdomen than chest; midbody scale rows across belly between ventrolateral folds 31;
scales on throat minute, granular, grading into larger scales on chest. Precloacal pores ina single series
of 9 with a narrow poreless scale separating two continuous rows of 4 (left) and 5 (right) pores; eleven
femoral pores in a continuous row on each thigh, separated by 9 scales from distalmost precloacal
pores; no precloacal groove. Scales on palm and sole smooth, flattened; scales on dorsal aspects of
hindlimbs flattened, similar to dorsal scales, with larger, conical tubercles interspersed among smaller
scales. Dorsal scales of forelimbs imbricate, without tubercles; scales of forearms heterogeneous with
scattered tubercles, some keeled.

Fore and hindlimbs relatively short, stout; forearm short (ForeaL/SVL ratio 0.16); tibia short
(CrusL/SVL ratio 0.18); digits long, strongly inflected at each joint, all bearing robust, recurved
claws; subdigital lamellae widened beneath basal phalanx; lamellae from first proximal scansor
greater than twice largest palm scale to first interphalangeal joint: 4-5-6-6-6 (manus) and 5-6-7-7-8
(pes); lamellae from first interphalangeal joint to toe tip, not including ventral claw sheath:
8-10-12-12-12 (manus) and 9-11-13-13-13 (pes); interdigital webbing present but weakly developed.
Relative length of digits (manus; measurements in mm in parentheses): III (10.0) > IV (9.8) > I (9.1)
> V (8.4) > I (6.6); (pes): V (12.0) > IV (11.2) > III (10.5) > II (8.7) > 1 (5.2).

82 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES
Volume 53, No. 7

Original tail long (TailL/SVL ratio 1.27), slender, slightly dorso-ventrally compressed. Scales
arranged in regular whorls; dorsal scales rectangular. Six circumferential rows per whorl; each whorl
with one row of 9 enlarged keeled tubercles, each tubercle separated from next by 0—2 smaller scales;
caudal tubercles reduced or absent on distal 2/3 of tail; posteriormost caudal scales narrow and elon-
gate. Subcaudal scales larger with a single median series of enlarged plates. Prominent cloacal spurs
with 3 enlarged scales on each side of tail base.

Osteology. Frontal bone moderately wide. Parietal bones paired. Premaxillary teeth loci 11, ap-
proximately 48 teeth on each maxillary bone, 54 on each dentary. Phalangeal formulae 2-3-4-5-3 for
manus and 2-3-4-5-4 for pes. Presacral vertebrae 26, including 3 anterior cervical (without ribs), 1
lumbar, and 2 sacral vertebrae; 5 pygal and 38 post pygal caudal in the complete original tail. One pair
of robust, crescentic cloacal bones present, at level of second pygal vertebra. (cloacal bones lacking 1n
female paratypes). Endolymphatic sacs not enlarged extracranially.

Coloration (in preservative). Base color a medium brown marked by numerous chocolate brown
patches, each with a dark brown border and a more peripheral, thin white line. Each marking bolder
and more well-defined posteriorly and more diffuse anteriorly. Head with light lines forming a reticu-
lated network between dark blotches. Occiput with three brown blotches (distinct in holotype but par-
tially or entirely fused in some paratypes). A series of smaller, more-or-less symmetrical dark
markings on top of head. Canthal region crossed by a pale, narrow transverse bar (faded in larger indi-
viduals, including holotype). A brown streak, bordered by cream white, running from behind orbit
onto neck; 6—8 (usually 7, 7 in holotype) paired markings on dorsum from nape to sacrum, left and
right sides often slightly out of phase. Markings on nape and sacrum generally oval, those between
axilla and groin partly bifurcate posteriorly. A series of more diffuse brown markings on lateral sur-
faces. Limbs more-or-less strongly barred, with alternating light and dark markings extending on to
digits. Tail base with elongate pair of dark marks similar to those on sacrum. Remainder of tail with al-
ternating elongate dark bands and shorter pale interspaces; each interspace with a dark center; 12 dark
bands, including tail tip, on original tail of holotype. Tail patterning extends on to venter, although
bands less clearly demarcated. Venter cream white with scattered light pigment on most surfaces
(males only). Palms and soles of feet dark in all specimens.

Color in life similar to that in preservative but with a slight yellowish cast to the white markings of
the head and body (Fig. 5).

VARIATION. — Comparative mensural data for the holotype and paratypes are presented in Table
2. Paratypes similar to holotype in most respects except as noted. Scale rows between ventrolateral
folds at midbody 27—32. Rows of dorsal tubercles at midbody 19-22. Precloacal pores 9—11, femoral
pores 12 oneach thigh in adult male paratypes (CAS 215474, 215661, USNM 548142). Females lack-
ing precloacal and femoral pores and without enlarged or dimpled scales in this region; cloacal spurs
present but weakly developed. CAS 210212, an adult female, contains two well-developed, shelled
eggs, each with a maximum diameter of 12.6 mm. Juveniles (CAS 210215—19, 215593, USNM
548141, 548143) with incompletely ossified long bones. Tail broken in 3rd post-pygal vertebra in
CAS 215661, 7th in USNM 548142, 18th in CAS 210215and 32nd in CAS 210212. All specimens
with original tails with 38 post-pygal vertebrae. Color patterns varies in number of paired dark dorsal
markings from 6 (CAS 210212) to 8 (CAS 210215, 215474), with all remaining specimens with seven
pairs. Each pair of markings may be highly symmetrical (e.g., CAS 210216, USNM 548141) or
largely out of phase (e.g., CAS 210217). Occipital markings distinct, as in holotype, or partially (CAS
210218, 215593, USNM 548142) or fully (CAS 210213, 210215, 210219, 215661, USNM 548143)
fused. No significant differences in any features were noted among the additional 27 specimens exam-
ined.

COMPARISONS. — Cyrtodactylus slowinskii may be distinguished from all congeners on the ba-
sis of its relatively large size (to 108 mm SVL), long digits and tail, absence of precloacal groove, se-
ries of 9-11 precloacal pores separated from a series of 11—12 femoral pores on each side, enlarged

BAUER: NEW SPECIES OF CYRTODACTYLUS 83

FiGureE 5. Living specimen of Cyrtodactylus slowinskii, sp. nov. (CAS 215483) from Alaungdaw Kathapa National Park,
Myanmar. Photo by Hla Tun.

subcaudal plates, and pattern of 6—8 pairs of more-or-less well-defined, dark dorsal blotches between
the nape and sacrum. Cyrtodactylus slowinskii may be distinguished from the following species by the
absence of a precloacal groove: C. annulatus (Taylor, 1915), C. cavernicolus (Inger and King, 1961),
C. fumosus (Miller, 1895), C. marmoratus (Gray, 1831), C. papuensis (Brongersma, 1934), C.
philippinicus (Steindachner, 1867), C. pubisulcus Inger, 1958, C. pulchellus Gray, 1827, C. rubidus
(Blyth, 1860), C. sadleiri Wells and Wellington, 1984; from the following species by the presence of
precloacal pores: C. jellesmae (Boulenger, 1897), C. laevigatus Darevsky, 1964, C. paradoxus
(Darevsky and Szczerbak, 1997), C. sermowaiensis (de Rooij, 1915), and most members of the subge-
nus Geckoella (C. collegalensis [Beddome, 1870], C. deccanensis [Ginther, 1864], C. jevporensis
[Beddome, 1877], C. nebulosus [Beddome, 1870], and C. yakhuna [Deraniyagala, 1945]); from the
following species by the presence of femoral pores: C. adleri Das, 1997, C. angularis (Smith, 1921),
C. brevidactylus Bauer, 2002 (this paper), C. elok Dring, 1979, C. fraenatus (Giinther, 1864), C.
ingeri Hikida, 1990, C. irianjayaensis Rosler, 2001, C. khasiensis (Jerdon, 1870), C. lateralis
(Werner, 1896), C. malayanus (de Rooij, 1915), C. matsuii Hikida, 1990, C. papilionoides Ulber,
1993, C. peguensis (Boulenger, 1893), C. quadrivirgatus Taylor, 1962, C. sworderi (Smith, 1925), C.
yoshii Hikida, 1990, and C. (G.) triedrus (Ginther, 1864); from C. biordinis Brown and McCoy, 1980
by having only a single row of femoral pores; from the following species by the presence ofa diastema
between precloacal and femoral pores: C. abrae Wells, 2002, C. derongo Brown and Parker, 1973, C.
Jarujini Ulber, 1993, C. loriae (Boulenger, 1898), C. louisiadensis (de Vis, 1892), C. novaeguineae
(Schlegel, 1844), C. tiomanensis Das and Lim, 2000, C. tuberculatus (Lucas and Frost, 1900), and C.
variegatus (Blyth, 1859); and from the following species by the presence of enlarged median
subcaudal plates: C. agusanensis (Taylor, 1915), C. brevipalmatus (Smith, 1923), C. gubernatorius
(Annandale, 1913), and C. wetariensis (Dunn, 1927). Of those species sharing with C. s/owinskii sim-
ilar precloacal and femoral pores and enlarged subcaudal plates, C. baluensis (Mocquard, 1890), C.
condorensis (Smith, 1920), C. consobrinoides (Annandale, 1905), C. consobrinus (Peters, 1871), C.
darmandvillei (Weber, 1890), C. feae (Boulenger, 1893), C. interdigitalis Ulber, 1993, C.
intermedius Smith, 1917, C. malcolmsmithi (Constable, 1949), C. mimikanus (Boulenger, 1914), C.
oldhami (Theobald, 1876), and C. redimiculus King, 1962 may be differentiated by their banded,
striped, or spotted dorsal patterns, and all except C. consobrinus and C. mimikanus are much smaller
(maximum SVL < 86 mm; Das 1997) than the new species. Cyrtodactylus irregularis (Smith, 1921) is

PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES

Volume 53, No. 7

NG
0'l
UES
Gi
69
O'S
8c
8°9
611
hgh)
L9G
(Qu
Br 69
cll
68
EL)

a[RUUdy

(Gia
OC
8°8
c8
Si)
9°8
hy)
oul)
C0C
V'8T
o Sv
18
96
0°61
VSI

S901

oyeul

II
60
CS
LY
69
LY
Sie
69
On
Vs
SC
vv
BINGz
Saul
v6
eso

d]BUId]

CE
fe I
SL
tee
SOI
VL
8°9
oTI
9°61
8LC
Cty
$6
qc Cl
O81
17

€ 10l

oyeul

SVO

ENC ve
Det 61
eS: ie)
LY SL
69 (NG)!
oS 6'8
(1y (G®)
69 vCl
OTI 10c
iS) (4 SOC
TSG L9V
7 16
BO'CL qs Cl
LOl 8S6l
C6 SSI
O19 8 LOI

oyeuloy oyeul

SVO SVO

ale
vl
£9
vs
SL
8s
Gs
68
6 vl
6 07
OSE
(ee)
B88
OI
VAG
8°6L
oyeul

SV)

SHE
6 |
g°9
6S
8
89
vs
16
Isl
8°0C
OLE
Lo)
®6'b6
vl
W) 6)
8°6L

o[PUloy

SVO

ie
81
59
v9
9'6
89
6S
06
eo
6 CC
v9E
78
80°98
VSI
Oe!
€€8
oy eu

SVO

Ye} =! (|

WMm~ MH

0°
3"
0°
9°
i:
z
.
:

Mm OM

I
6 61
O'CE
vs

®0'r6
6 Cl
vol
ofl,

ayeuldy

SVO

OC
ia
6S
61
Mi
61
CV
VL
Gl
VLI
0°6C
LS
q9'cr
GI
Cc Ol
9199.
oyRUldy

SV)

Se ve
vC 81
9°8 SE
L’8 Ons
sil vol
L8 coal
OL II)
OCI SOI
L‘0C V8
6 8C v9C
eLy COV
8°6 v8
BCOEl 40801
681 Al
091 esl
L’Yyol L96

9] PUloy 9] PUloy

EVcVemCyLOVcmmbiLSvs S LOOSTGmsEOSSIC PLYSIC™ 6IZOLC SIZOIC ZIZOIZ SIZOIZ” SIZOIS €17OIt Zicolz
WNSO WNSN WNSN

SVO SVO

oe
OC
$8
08
901
Pye
2)
eo}
lol
LIC
Lv
9°8
80'SCI
eal
aa
8°86
oyeul

vICOLT

SVO

JRUIOU]
Led
QIO1OU]
oAqIeN
aAqus
IeqoAq
dado
HP®2°H
MPP®°H
‘TP®°H
qyunsy
MTEL
TRL
Ysnip
[eo104

TAS
XO

‘poyesroudsaI = 9 “Udyo.1g = q ‘[RUITIIO =v Tel -sodAqesed ore suowiisads Joyo ‘adAjojoy ayy SI PI ZOIZ sy) OJON
“WU UT SJUSWIDINSBOUT []B “SpOyjJo|] Pur SyeLIo}B\] Ul se SUOTIeIADIQGY “AOU “ds “12ysUIMO]S SN]AJIDPOJAAD JO Saisas 9dAq dy} OJ vIep [eINSUd\ ‘7 ATAV

BAUER: NEW SPECIES OF CYRTODACTYLUS 85

smaller (maximum SVL 79 mm fide Smith 1935) and has a much larger number of midventral scales
(41-46 vs. 27-32).

DISTRIBUTION. — Cyrtodactylus slowinskii has thus far been recorded only from Alaungdaw
Kathapa National Park in the Sagaing Division of Myanmar, where it appears to be both common and
widespread. It is probable that the species is more widely distributed along the eastern foothills of the
Rakhine Yoma (Arakan Yoma).

DISCUSSION

The affinities of Cyrtodactylus brevidactylus are unclear. Precloacal and femoral pore character-
istics, although generally specifically stable, vary widely in the genus and are not obvious markers of
interspecific affinities. Cyrtodactylus brevidactylus bears little resemblance to any of its congeners in
respect to its rugose scalation and body shape. Likewise, its striking short digits are unique within the
genus, although somewhat similar in proportion to those of some members of the Indian-Sri Lankan
subgenus Geckoella, with which C. brevidactylus also shares a similar general habitus. The species is
also very similar in body proportion to members of the Eublepharidae, although it lacks moveable
eyelids and the osteological features (e.g., unpaired parietal bone) diagnostic of these gekkotans.
Among regional geckos, its color pattern is most nearly approached by C. angularis and C.
papilionoides of Thailand. These species share the small number of large, dark, dorsal blotches, but
differ in the pattern of the dorsum of the head (see Chan-Ard et al. 1999, for color photographs of these
and other Thai and Malaysian species of Cyrtodactylus).

Cyrtodactylus slowinskii is more typical of the genus in body form and superficially resembles
other moderately large members of the genus. Its pattern of precloacal and femoral pores is shared
with a number of geographically proximate congeners and it is probably allied to morphologically
similar species such as C. consobrinus, although in the absence of a robust hypothesis of relationships,
specific affinities remain conjectural at best.

Preliminary examination of additional collections from Myanmar suggests that at least two addi-
tional undescribed Cyrtodactylus are present, bringing the national total to twelve. The diversity of the
herpetofauna of Myanmar has long been regarded as lower than that of surrounding areas. In large part
this is due to the limited collecting activity in the country prior to the last several years (Inger 1999;
Slowinski and Wiister 2000). Preliminary results of the on-going Myanmar Herpetofaunal Survey
suggest that Myanmar is herpetogeographically complex, supporting taxa of both Indian and south-
east Asian affinities, as well as endemics. Slowinski and Wiister (2000) described the Burmese spit-
ting cobra (Naja mandalayensis) from the central dry zone of Myanmar and proposed that this region,
lying between the Rakhine Yoma (Arakan Yoma) in the west and the extensive montane areas east of
the Sittaung River (Fig. 3), would prove to be a significant area of endemism. The new Cyrtodactylus
described herein support this proposition and the apparent restriction of C. brevidactylus to Mt. Popa
further suggests that isolated physiographic features within the central dry zone may harbor endemics
of their own.

ACKNOWLEDGMENTS

I thank George Zug and the late Joseph B. Slowinski for inviting me to examine the geckos de-
scribed herein. Specimens for this report were obtained by the Myanmar Herpetofaunal Survey, a
joint program of the Myanmar Nature and Wildlife Conservation Division, Forest Department; the
California Academy of Sciences; and the Smithsonian’s National Museum of Natural History with
support from the National Science Foundation (Grant DEB 9971861). In particular I thank the 2000
Myanmar Herpetofaunal Survey team: U Htun Win, Daw Thin Thin, U Hla Tun, U Sai Wunna Kyi, U
San Lwin Oo, U Kyi Soe Lwin, and U Awan Khwi Shein. Robert C. Drewes and Jens V. Vindum
(CAS), George Zug, Robert Wilson and Ken Tighe (USNM), and James Hanken and José Rosado

86 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES
Volume 53, No. 7

(MCZ) provided access to specimens in their care. Michelle Koo kindly prepared the map. The manu-
script benefited from the comments of George Zug, Indraneil Das and Jens Vindum.

LITERATURE CITED

BAUER, A. M. AND K. HENLE. 1994. Familia Gekkonidae (Reptilia, Sauria). Part | Australia and Oceania. Das
Tierreich 109 (part):i—xin1 + 1-309.
CHAN-ARD, T., W. GROSSMANN, A. GUMPRECHT, AND K-D. SCHULZ. 1999. Amphibians and reptiles of peninsu-
lar Malaysia and Thailand, an illustrated checklist. Bushmaster Publications, Wuerselen, Germany.
DAREVSKY, I. S. AND N. N. SZCZERBAK. 1997. A new gecko of the genus Gonydactylus (Sauria: Gekkonidae)
with a key to the species from Vietnam. Asiatic Herpetol. Res. 7:19—22.

Das, I. 1997. A new species of Cyrtodactylus from the Nicobar Islands, India. J. Herpetol. 31:375—382.

Das, I. AND L. J. Lim. 2000. A new species of Cyrtodactylus (Sauria: Gekkonidae) from Pulau Tioman, Malaysia.
Raffles Bull. Zool. 48:223—231.

HIKIDA, T. 1990. Bornean gekkonid lizards of the genus Cyrtodactvlus (Lacertilia: Gekkonidae) with descrip-
tions of three new species. Japan. J. Herpetol. 13:91—107.

HUNDLEY, H. G. 1964 et seq. Check list of reptiles of Burma. Burmese Forestry Department internal document
[mimeograph], pp. 1-111, 1-28 [1964]; also Supplement I. Check list of reptiles of Burma, pp. 1—7 [ca. 1965].

INGER, R. F. 1999. Distribution of amphibians in southern Asia and adjacent islands. Pp. 445-482 in Patterns of
distribution of amphibians, a global perspective, W. E. Duellman, ed. Johns Hopkins University Press, Bal-
timore.

KHAN, M.S. AND H. ROSLER. 1999. Redescription and generic redesignation of the Ladakhian gecko
Gymnodactylus stoliczkai Steindachner, 1969 [sic]. Asiatic Herpetol. Res. 8:60—68.

KLUGE, A. G. 1985. Notes on gecko nomenclature (Sauria: Gekkonidae). Zool. Meded., Leiden 59:95—100.

—. 1991. Checklist of gekkonoid lizards. Smithson. Herpetol. Inform. Serv. (85):1—35.

—. 1993. Gekkonoid lizard taxonomy. International Gecko Society, San Diego.

. 2001. Gekkotan lizard taxonomy. Hamadryad 26:1—209.

MEES, G. F. 1987. On the generic names Gonydactylus Kuhl and van Hasselt, 1822, Goniodactylus Schlegel,
1826, and Gonydactylus Wagler, 1830 (Reptilia: Sauria: Gekkonidae). Zool. Meded., Leiden 61:137—140.

ROSLER, H. 2000. Kommentierte Liste der rezent, subrezent und fossil bekannten Geckotaxa (Reptilia:
Gekkonomorpha). Gekkota 2:28—153.

. 2001. Eine neue groiwichsige Cyrtodactylus-Art von Neuguinea (Reptilia: Sauria: Gekkonidae). Zool.
Abh. Staatl. Mus. F. Tierk. Dresden 51:61—71.

SLOWINSKI, J. B. AND W. WUSTER. 2000. A new cobra (Elapidae: Naja) from Myanmar (Burma). Herpetologica
56:257-270.

SMITH, M. A. 1935. The fauna of British India, including Ceylon and Burma. Reptilia and Amphibia. Vol. II —
Sauria. Taylor and Francis, London.

SZCZERBAK, N. N. AND M. L. GOLUBEV. 1977. Systematics of the Palearctic geckos (genera Gymnodactylus,
Bunopus, Alsophylax) Trudy Zool. Inst. Akad. Nauk USSR 74:120—133. [In Russian]

. 1984. On generic assignement of the Palearctic Cyrtodactylus lizard species (Reptilia, Gekkonidae).

Vestnik Zoologi 2:50—S6. [In Russian]

. 1986. Gecko fauna of the U. S. S. R. and contiguous regions. Naukova Dumka, Kiev.

ULBER, T. 1993. Bemerkungen tiber cyrtodactyline Geckos aus Thailand nebst Beschreibungen von zwei neuen
Arten (Reptilia: Gekkonidae). Mitt. Zool. Mus. Berlin 69:187—200.

UNDERWOOD, G. 1954. On the classification and evolution of geckos. Proc. Zool. Soc. London 124:469-492.

WELLS, R. W. 2002. Taxonomic notes on the genus Cyrtodactylus (Reptilia: Gekkonidae) in Australia. Austral.
Biodiv. Rec. (3):1—8.

-
amt bel meeren

b, a S
z = ary ds Wy ¢ NF Mattes ®nal inte
oh ina) Le Ge & it Creal Ey ‘ine wi) | ee eid

= =a

PF) yer in Cie

2
‘ - = Q
’ —— 44 . flavin (SS > Pre i ay
- *
= a
4 j pts
ams Wy 0
uw aa
te 1 1. =
° L wie, MPs s a4 i@-* '
ei 1 * : : ’ :
te we eek

PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES

Volume 53, No. 8, pp. 87—93, | pl., 1 table. July 1, 2002

Parapercis lata, a New Species of Sandperch (Perciformes:
Pinguipedidae) from the Central Pacific

by
John E. Randall
Bishop Museum, 1525 Bernice St., Honolulu, Hawaii 96817-2704
| JUL 0 8 2007
and

John E. McCosker ;
California Academy of Sciences, Golden Gate Park, €alifornia 94118 _

Parapercis lata is described from 12 specimens from the Line Islands and one from the
Phoenix Islands collected over sand or rubbJe substrata in 6—55 m in both lagoon and
outer-reef areas. It is distinct in having the anterior body broader than its depth, three
pairs of canine teeth at the front of the lower jaw, no palatine teeth, a short spinous portion
of the dorsal fin with V spines, the last connected by membrane directly across to first soft
ray, 59-60 lateral-line scales, and a unique color pattern of eight narrow black bars on the
body, the middle six bars branching dorsally to a Y-shape.

The Pinguipedidae, established as the current name to replace the Parapercidae and Mugiloididae
of the older literature (Rosa and Rosa 1987), consists of six genera. The four that are universally rec-
ognized are Kochichthys, Parapercis, Pinguipes, and Prolatilus. We include the monotypic freshwa-
ter genus Cheimarrichthys from New Zealand, following McDowell (1973), and Pseudopercis,
demonstrated as valid for the family by Rosa and Rosa (1997). The largest genus is Parapercis, all
species of which are found in marine habitats, mostly in the Indo-Pacific region, with a single south-
eastern Pacific species (McCosker 1971). Cantwell (1964) first revised Parapercis, recognizing 27
species. Ina brief review of the genus, Randall (1984) raised the total to 40 species. Manilo (1990) de-
scribed P. ventromaculata as new from the Maldive Islands, but this is ajunior synonym of P. signatus
Randall, 1984, also from the Maldives. Six more species have been described to date (Anderson 1992;
Randall and Francis 1993; Randall, in press), but others await description.

The first author has long planned a second revision of this genus, but this has been repeatedly
postponed. The species described herein was first collected in the Line Islands in 1968; its description
was intentionally delayed for the revision. However, after finding a specimen in the collection of the
California Academy of Sciences from the Phoenix Islands, it was decided to describe the species so
the name will be available for a large manuscript in progress by the first author on the reef and shore
fishes of the South Pacific.

The species of Parapercis, popularly known as sandperches, are generally found on sedimentary
or rubble substrata, often near coral reefs. They are usually seen at rest on the bottom, propped on their
pelvic fins, but they often move short distances as they forage for food—chiefly small benthic crusta-
ceans and demersal zooplankton. Several authors have noted the sexual dichromatism of species of
the genus, and protogynous hermaphroditism has been demonstrated for some; also males are known
to be haremic (Randall 1984; Nakazono etal. 1985; Clark et al. 1991; T. J. Donaldson, pers. comm.).

88 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES
Volume 53, No. 8

MATERIALS AND METHODS

Type specimens of the new Parapercis have been deposited at the following institutions: Austra-
lian Museum, Sydney (AMS); Bernice P. Bishop Museum, Honolulu (BPBM); California Academy
of Sciences, San Francisco (CAS); Museum National d’Histoire Naturelle, Paris (MNHN); and the
U. S. National Museum of Natural History, Washington, D. C. (USNM).

Lengths of specimens are given as standard length (SL), the distance from the front of the upper
lip to the base of the caudal fin (posterior end of the hypural plate). Body depth is measured vertically
from the origin of the anal fin; body width is taken just posterior to the gill opening. Head length is
measured from the front of the upper lip to the posterior end of the opercular membrane, and snout
length from the same anterior point to the fleshy edge of the orbit. Orbit diameter is the greatest fleshy
diameter, and interorbital width the least fleshy width. Upper-jaw length is taken from the front of the
upper lip to the fleshy end of the maxilla. Dorsal and anal spines and soft rays are measured from the
point they depart from the contour of the body.

Counts of lateral-line scales are made to the base of the caudal fin, hence do not include the series
of six or seven small pored scales on the base of the fin.

In the description of the new species, data in parentheses apply to paratypes. Proportional mea-
surements in the text are rounded to the nearest 0.05.

SPECIES DESCRIPTION

Parapercis lata n. sp.
Plate 1, Figs. a—d; Table 1

MATERIAL EXAMINED. — HOLOTYPE: BPBM 28063, male, 212.0 mm, Line Islands, Tabuaeran
Atoll (Fanning Island), about one-half mile southeast of entrance to English Harbor, ocean reef slope,
sand and rubble, 36.5 m, net, J. L. Earle, 3 April 1981. PARATYPES: BPBM 7575, 186.0 mm,
Tabuaeran, lagoon end of English Harbor off settlement, sand bottom with isolated coral heads, spear,
J. E. Randall, 29 October 1968; AMS 1.141173-001, 128.6 mm, same locality, 6 m, spear, J. E.
Randall, 31 October 1968; CAS 42102, 174.0 mm, Phoenix Islands, Canton Atoll, Shark Alley, spear,
D. Diener, 15 August 1978; BPBM 28066, 2: 143.2-164.5 mm, Line Islands, Tabuaeran, ocean side
offcable station, sand channel with coral heads, soft coral, and rubble, 20-21 m, spear and quinaldine,
J. E. Randall, 4 April 1981; BPBM 30657, 74.8 mm, Line Islands, Kiritimati Atoll (Christmas Is-
land), net, D. Wilder, July 1985; USNM 367667, 145.4 mm, Kiritimati, NE side off Captain Cook Ho-
tel, outer reef, rubble and sand, 55m, spear, J. E. Randall, 29 July 1987; BPBM 31908, 3:
97.3-178.8 mm, Kiritimati, off London, north of Bridges Point, rubble, sand, and patch reefs,
12-13 m, spear, J. E. Randall, 2 August 1987; MNHN 2002-0138 , 132.0 mm, Kiritimati, west side,
rotenone, 6-12 m, D. R. Robertson, September 1996.

DIAGNOSIS. — Dorsal rays V,21; anal rays I,17; pectoral rays 18-19 (usually 18); lateral-line
scales 59-60; predorsal scales about 15; scales on cheek small and cycloid; gill rakers 6—7 + 10-12;
three pairs of canine teeth anteriorly in lower jaw, the medial pair small, the lateral largest; palatine
teeth absent; no serrae or spines on edges of opercular bones except a single stout spine on opercle;
body elongate, the depth 5.7—6.2 in SL; body cylindrical anteriorly, the width slightly greater than
depth; third or fourth dorsal spines longest; membrane of fifth dorsal spine connected directly across
from spine tip to first soft ray; caudal fin slightly rounded, the upper corner projecting as a short,
broad-based filament in adults; whitish with eight narrow dark bars on body, all but first and last
broadly expanded dorsally to a Y-shape; largest specimen, 212 mm SL.

DESCRIPTION. — Dorsal rays V,21; anal rays I,17; pectoral rays 18 (18, three of 11 paratypes
with 19); pelvic rays I,5; principal caudal rays 16, the median 14; branched; upper procurrent caudal

RANDALL AND McCOSKER: A NEW SPECIES OF SANDPERCH

PLATE |

A. Holotype of Parapercis lata, BPBM 28063, 212 mm SL, Tabuaeran, Line Islands.
B. Female of Parapercis lata, BPBM 28066, 143.2 mm SL, Tabuaeran, Line Islands.
C. Subadult of Parapercis lata, BPBM 30657, 74.8 mm SL, Kiritimati, Line Islands.
D. Underwater photograph of male of Parapercis lata, about 225 mm TL, Kiritimati,

Line Islands.

90 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES
Volume 53, No. 8

rays 9 (9-10), the last segmented; lower procurrent caudal rays 8 (8-9), the last segmented; lat-
eral-line scales 59 (59-60) + 6—7 progressively smaller pored scales on caudal-fin base; scales above
lateral line to origin of dorsal fin 9; scales below lateral line to origin of anal fin 14; circumpeduncular
scales 30; predorsal scales about 15; gill rakers 6 + 10 (6—7 + 10-12); pseudobranchial filaments 32
(14-28, increasing with growth); branchiostegal rays 6; vertebrae 30.

Body elongate, the depth 6.2 (5.7—7.2) in SL; body width just behind gill opening greater than
depth, the width 5.8 (5.45—7.0) in SL; head length 3.35 (3.25—3.4) in SL; snout pointed, its length 2.7
(2.72.95) in head length, its anterior end rounded in dorsal view; orbit diameter 6.1 (4.4—5.9) in head;
interorbital space slightly concave, its width 6.65 (6.6—9.6) in head; caudal-peduncle depth 3.0
(2.9-3.3) in head; caudal-peduncle length 3.5 (3.45—3.9) in head.

Mouth slightly oblique, forming an angle of about 20° to horizontal axis of head and body, the
lower Jaw projecting; upper jaw extending to or slightly posterior to a vertical at front edge of orbit, the
upper-jaw length 2.6 (2.5—2.9) in head; front of upper jaw with 6—7 pairs of incurved canine teeth, the
most posterior usually the largest, followed by a row of slender incurved conical teeth (24 on side of
jaw of holotype); a broad band of villiform teeth immediately behind anterior canines, narrowing to a
single row posteriorly in jaw; front of lower jaw with 3 pairs of incurved canine teeth, progressively
larger laterally, the third pair larger than largest canine of upper jaw; a broad band of villiform teeth
immediately behind anterior canines, narrowing to a single row of slender conical teeth posterior to
canines that enlarge to a series of 4 or 5 recurved canines on midside of jaw; a narrow band of small
conical teeth in a broad arc on vomer; no teeth on palatines. Inner surface of lips with ridges of fleshy
papillae that extend between anterior canines. Tongue narrowly triangular witha slightly rounded tip.

Gill membranes united, not attached to isthmus, with a broad free fold across. Gill rakers short
and stout, less than one-fourth length of longest gill filaments on first gill arch. Anterior nostril small
with a rim and posterior flap (not reaching posterior nostril when laid back), in front of middle of eye a
distance of three-fourths eye diameter in holotype; posterior nostril about twice as large, with a slight
rim, dorsoposterior to anterior nostril, the internarial distance one-fifth orbit diameter in holotype.

Opercle with a strong horizontal spine in line with front of upper lip; no other spines on head, and
no serrae on edges of opercular bones; preopercle broadly rounded, its free posterior margin ending at
level of lower edge of orbit; opercular membrane broadest ventrally where it joins branchiostegal
membrane.

Lateral line continuous, approximately following contour of back; lateralis system of head well
developed, with numerous sensory pores, the four on each side of mandible and one symphyseal the
most prominent. Scales on body ctenoid, those on side of body with about 50 cteni on holotype and
about 25 on smallest paratype; scales on opercle and cheek very small and cycloid, partially embed-
ded on opercle and fully embedded on cheek, those on cheek reaching slightly anterior to corner of
mouth; no scales on dorsal, anal, or pelvic fins; small scales on about basal one-fourth to one-fifth of
pectoral fins; very small scales extending about three-fourths length of caudal fin.

Origin of dorsal fin over fourth lateral-line scale, the predorsal length 3.2 (3.05—3.2) in SL;
spinous portion of dorsal fin low, the spines progressively shorter with growth; first dorsal spine 15.5
(11.5—15.0) in head length; third and fourth dorsal spines subequal, the longest 6.35 (4.4—6.5) in head;
fifth dorsal spine 9.1 (7.35—9.55) in head; membrane from fifth dorsal spine attached directly across to
first soft dorsal ray; dorsal soft rays except first two and last subequal, the longest 2.1 (2.05—2.6) in
head; membranes of anterior soft portion of dorsal fin incised about one-half length of rays, grading to
one-third posteriorly; origin of anal fin below base of fifth dorsal soft ray, the preanal length 2.05
(2.05—2.1) in SL; anal-fin spine very slender and closely applied to first soft ray, 6.25 (5.35—7.05) in
head; 15th and 16th anal soft rays longest, 2.65 (2.4—2.75) in head; caudal fin slightly rounded, the
first two branched rays slightly prolonged, forming a broad-based filament; caudal-fin length 5.0
(4.8-5.05) in SL; pectoral fins rounded, the middle rays longest, 5.75 (5.3—5.85) in SL; origin of pel-
vic fins slightly anterior to base of pectoral fins, the prepelvic length 3.75 (3.5—3.8) in SL; pelvic spine

RANDALL AND McCOSKER: A NEW SPECIES OF SANDPERCH 9]

TABLE |. Proportional measurements of type specimens of Parapercis lata expresses as percentages of
the standard length.

Holotype Paratypes

BPBM BPBM BPBM BPBM BPBM BPBM BPBM_ BPBM
28063 30657 = 31908 += 331908 »3=—s 28066 )=— 28066) = 31908 TIS

Sex male female female female female male male male
Standard length (mm) 212.0 74.8 O73 125.0 143.2 164.5 178.8 186.0
Body depth 16.2 13.9 14.5 17-5 15.4 [Sal 16.8 16.5
Body width WAS) 14.3 16.5 18.4 16.8 16.4 179 17.4
Head length 29.9 29.4 DoS 30.8 30.3 30.0 29.6 30.0
Snout length Me 10.1 10.0 St leis 10.6 10.6 10.6
Orbit diameter 4.9 6.7 6.2 6.1 5.6 5.4 Spal Spl
Interorbital width 4.5 el 3.4 3:2 4.2 3.8 4.5 4.5
Upper-jaw length 11.4 10.8 Li2 10.7 122 12.0 113 MES
Caudal-peduncle depth poe 9:1 9.0 10.5 9.4 9.1 10.1 10.0
Caudal-peduncle length 8.5 7.6 8.0 8.0 8.2 8.6 8.2 8.7
Predorsal length B15 32.0 31,3 Sai 32.0 eyes) Sie, ail
Preanal length 47.5 48.3 48.1 49.1 47.3 48.8 48.2 47.6
Prepelvic length 2647, 27.0 26.7 29.6 26.5 27.0 27a 27.4
Dorsal-fin base 60.6 61.7 60.5 61.0 60.3 60.4 61.2 60.4
First dorsal spine 19 2.6 25 23 2.4 pe) 22 2.0
Longest dorsal spine 4.7 6.7 29 5.6 5.4 Sel 4.7 4.6
Fifth dorsal spine 3.3 4.0 3.3 BS) 3.4 3:2 3.1 3.4
Longest dorsal ray We 12.4 In (MAS) 11.8 11.6 lull5 i)
Anal-fin base 43.2 44.7 43.4 44.2 43.0 42.7 43.7 43.1
Anal-fin spine 4.8 3/45) a5 4.8 4.3 4.6 4.9 4.7
Longest anal ray a Gs} 12.4 11.6 P2 ates) A th2 1LS
Caudal-fin length 20.0 2037 19.8 20.8 20.2 20.1 20.4 20.3
Pectoral-fin length 17.4 18.8 18.2 18.4 18.4 18.1 17.1 eZ
Pelvic-spine length 7.4 8.4 8.1 8.3 8.4 8.5 7.8 7.6
Pelvic-fin length 18.6 2371 22.6 22.0 21.6 197 19.0 18.8

very slender and closely applied to first soft ray, its length 4.05 (3.5—3.95) in head; fourth pelvic soft
ray longest, just reaching anus in holotype, but extending beyond origin of anal fin in smaller
paratypes, 5.4 (4.3—5.3) in SL.

Color of holotype, a male, in alcohol very pale brown (nearly white) with eight dark brown bars
less than half width of pale interspaces, the middle six of which bifurcate dorsally to form a broad
Y-shape; a brown bar containing small brown spots and short lines across occiput, angling down to
cross pectoral-fin base and chest; scales of pectoral base and chest bar and first five bars on body with
dark brown centers and pale edges thus giving a vertical linear effect within these bars; pale
interspaces on body with one to three faint irregular pale yellowish brown bars; a curved brown band
across top of snout from eye to eye; a dark brown line and two small dark brown spots extending poste-
riorly from eye in line with a curving dark brown line within brown bar of posterior part of head; an
oblique dark-edged band (yellow in life) from lower edge of orbit across operculum; last three dark
bars on body with two small dark brown spots; scattered dark brown dots dorsally on postorbital head
and anterodorsally on body from beneath pectoral fin to upper part of fifth dark bar on body; four dark
brown dots in interorbital space and a median one just before; a dark brown spot anteriorly on side of
upper lip; dorsal and anal fins translucent with pale yellowish rays, the dorsal with a row of small dark
brown spots at base, one on every other membrane, and a second row of fainter spots above on mem-

92 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES
Volume 53, No. 8

branes without a dark spot below, except posterior six membranes with an upper spot on each mem-
brane, these now darker; a dark brown dot at tip of each membrane of soft portion of dorsal and anal
fins; anal fin with four small dark brown spots posteriorly; caudal fin pale yellowish with five faint ir-
regular narrow brown bars containing dark brown dots; paired fins with translucent membranes and
pale yellowish brown rays. Color of holotype when fresh shown in Plate la.

The following color note was made of the 186-mm male paratype of BPBM 7575 when fresh:
pale olive, shading on sides and ventrally to white, with eight blackish bars on body and one posteri-
orly on head crossing to chest, the median six of these bars branched dorsally; bar on head and last
three bars on body with small black spots; white interspaces on body with one to three irregular light
olivaceous yellow bars; black dots on postorbital head and anterodorsal part of body; a diagonal
dark-edged olive bar passing posteroventrally from eye; a narrow black band curving on top of snout
just before interorbital space, passing horizonally through eye to upper end of preopercular margin; a
dotted black line across middle of interorbital space; fins pale, the dorsal and caudal with black dots;
dorsal fin with a pinkish margin (mainly ray tips); anal fin and middle of caudal fin with a narrow
white margin and broad pinkish submarginal zone.

The color of the 143.2-mm female paratype of BPBM 28066 is shown in Plate 1b, and that of the
74.8-mm subadult of BPBM 30657 in Plate 1c. The underwater photograph of Plate 1d provides the
life color of a male estimated at 225 mm total length.

Males can be distinguished from females by the oblique, dark-edged, yellow or olive band on the
cheek, more numerous black dots dorsally on the head and dorsoanterior part of the body, and the
more pointed membrane at the tips of the dorsal spines.

ETYMOLOGY. — This species is named Parapercis lata from the Latin for wide, in reference to
the width of the anterior part of the body, greater than the depth for all specimens examined.

REMARKS. — Parapercis lata is presently known only from the atolls of Tabuaeran and
Kiritimati in the Line Islands and Canton in the Phoenix Islands; it appears to be endemic to this part of
the Central Pacific. Because it is large and easily approached, it surely would be known from
well-collected island groups, such as the Marshall Islands, Samoa Islands, and the islands of French
Polynesia, if it were there.

Specimens were collected from sand or sand and rubble substrata in both lagoon and seaward reef
areas at depths of 6-55 m.

Of the known species of Parapercis, P. lata is closest to P. xanthozona (Bleeker) which occurs
from East Africa to the western Pacific, ranging east in Oceania only to Palau, New Caledonia, and
Fiji. The two species share the same meristic data, the same general morphology, dentition, and large
size. P. lata differs in having a shorter spinous portion of the dorsal fin, the longest spine less than
one-half the length of the longest soft ray, whereas in P. xanthozona the longest spine is from one-half
to two-thirds the length of the longest soft ray. The width of the body measured just behind the gill
opening is greater than the body depth in P. /ata, but it is equal to or less than the depth in P.
xanthozona. Also the two are very different in color, P. xanthozona having a conspicuous midlateral
white stripe that extends to the end of the caudal fin, dark bars only below the white stripe, a large yel-
low spot at the base of the pectoral fins, and the male with a brownish yellow cheek crossed by oblique
dark-edged white lines.

The four largest type specimens are males, the rest are females. The gonad of the smallest male,
BPBM 28066, 164.5 mm SL, contains yellow inclusions which seem to be ova in the process of being
resorbed. This suggests that Parapercis lata is a protogynous hermaphrodite, as has been shown for a
few other species of the genus.

RANDALL AND McCOSKER: A NEW SPECIES OF SANDPERCH 93

ACKNOWLEDGMENTS

We thank John L. Earle, Douglas Diener, D. Ross Robertson, and David Wilder for collecting
specimens of the new Parapercis, Loreen R. O’Hara for X-rays, Alan Leviton for assistance with
Plate 1, and Tomio Iwamoto for reviewing the manuscript.

LITERATURE CITED

ANDERSON, M. E. 1992. A new sandperch Parapercis maritzi (Teleostei: Pinguipedidae) from South Africa. S.
Afr. J. Zool., 27(4):151—156.

CANTWELL, G. E. 1964. A revision of the genus Parapercis, family Mugiloididae. Pac. Sci. 18(3):239-280.

CLARK, E., M. POHL, AND J. RABIN. 1991. Spotted sandperch dynamics. Natl. Geogr. Res. and Explor.
7(2):138-155.

MANILO, L. G. 1990. New species of the genus Parapercis (Mugiloididae) from the region of the Maldive Is-
lands. Voprosy Ikhtiologii 30(6):1016—1019.

McCosker, J. E. 1971. A new species of Parapercis (Pisces: Mugiloididae) from the Juan Fernandez Islands.
Copeia 1971(4):628—686.

McDowaALL, R. M. 1973. Relationships and taxonomy of the New Zealand torrent fish, Cheimarrichthys forsteri
Haast (Pisces: Mugiloididae). J. Roy. Soc. New Zeal. 3(2):119—227.

NAKAZONO, A., H. NAKATANI, AND H. TSUKAHARA. 1985. Reproductive ecology of the Japanese reef fish,
Parapercis snyderi. Proc. Fifth Internatl. Coral Reef Congr., Tahiti 5:355—360.

RANDALL, J. E. 1984. Two new Indo-Pacific mugiloidid fishes of the genus Parapercis. Freshw. and Mar. Aquar.
7(12):41-49.

RANDALL, J. E. In press. Review of the sandperches of the Parapercis cylindrica complex (Perciformes:
Pinguipedidae), with description of two new species from the western Pacific. Occ. Pap. B. P. Bishop Mus.

RANDALL, J. E. AND M. P. FRANCIS. 1993. Parapercis colemani, a new pinguipedid fish from Norfolk Island,
south-western Pacific Ocean. New Zeal. J. Mar. and Freshw. Res. 27:209-214.

Rosa, I. L. AND R. S. ROSA. 1987. Pinguipes Cuvier and Valenciennes and Pinguipedidae Gunther, the valid
names for the fish taxa usually known as Mugiloides and Mugiloididae. Copeia 1987(4):1048—1051.

. 1997. Systematic revision of the South American species of Pinguipedidae (Teleostei, Trachinoidet).

Revta Bras. Zool. 14(4):845—865.

.
-

janet Oe CG0y pot hi hr) ate
fone! hen tome Sep tes, Vleet Trae |
stagnate diane

peels ne% Sning euler (A fi writ al ae O° <a horery pent
t jiugtinY wads crane ) sve)cr: ipitea et et cabin >

Yay i, @ ve tab lasy, ts 11) OP ef id at aN

a i

al?

Say alta PrivcrOSE EAT ahah - Meare
arrow es ae Si eee x! Ty yh e a it

sae MTL Vet tal cae iw sited wees iattnred sapoty Oy
jumpy: bewi 158 Mee iat) Rie ipsa bait sie anki’ i

eet, Ty" dase 7 . uve ytwiel ore

T tiug ; y
SWAY We a cant (ned Jed ved) eyes et) § Gan rl aires

, : bain es, ieohasa i
oy nee Gi oocmwd i eran os

= be) i
er ke Pe Lene meen en
ty oat! A ie

: , Ye ; vin ut ky
' “Ms ‘ : é a) A ey Ae a
log) One

PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES

Volume 53, No. 9, pp. 95-116, 3 figs. SEP 13 2002 August 22, 2002

Systematics of the Genus Monolena (Melastomataceae)
in Central America

by
Richard H. Warner

10401 Chesterwood Dr., Spotsylvania, Virginia 22553
email: omar(@va.prestige.net

Morphological, anatomical, and field studies indicate that Monolena (Melastomataceae) is
represented by eight species in Central America. Seven of these species are found in Pan-
ama: Monolena grandiloba, M. dressleri, M. multiflora, M. panamensis, M. morleyi, and M.
trichopoda, newly described and all endemic to Panama, and M. primuliflora, which ranges
from eastern Costa Rica to southern Peru. The eighth species, M. guatemalensis, is endemic
to Guatemala. Taxonomic accounts of these species are presented, along with notes on
anatomy, cytology and distribution.

The genus Monolena (Melastomataceae) consists of 14-16 species with an overall distributional
range from Guatemala to southern Peru and Acre, Brazil. The group is distinguished from other mem-
bers of the Melastomataceae by the combination of: (1) its herbaceous habit, in which the stem is re-
duced to a fleshy rhizome; (2) its l-sided pedunculate and bracteate cyme (or I-flowered scape); (3) its
3-locular ovary in an otherwise 5-merous flower; (4) its dimorphic androecium; (5) its anthers, each
of which has the connective ventrally extended between the thecae and the insertion of the filament
and further extended to form a large or small ventral appendage; and (6) its triquetrous capsular fruit.
Members of the genus are mostly epiphytic, growing on lower trunks and branches, buttresses,
stumps, logs and rocks, or sometimes on the ground.

TAXONOMIC HISTORY

The earliest description of Monolena was provided by J. J. Triana in Bentham and Hooker
(1867). Triana had collected several species of Monolena from western Colombia and had apparently
seen other collections from Peru, but he described no species and cited no specimens with the first ge-
neric description. Monolena primuliflora Hook.f. was the first species described, that being done in
1870.

A later publication from Triana (1871) described one species from western Colombia. Cogniaux
(1891) and Gleason (1930) each published a species from eastern Colombia, and Uribe (1960, 1971,
1979) described four more species from western Colombia. Prior to the present paper, Monolena
guatemalensis Donn. Sm. was the only species of Monolena described from Central American plants.
Monolena was treated in regional floras for Guatemala (Standley and Williams 1963), Panama
(Gleason 1958), Ecuador (Wurdack 1980), and Peru (MacBride 1941). The genus belongs to the New
World tribe Bertolonieae; that tribe could possibly be merged with the Old World Sonerileae
(Wurdack 1980). A revision of Monolena became necessary when many diverse new taxa were dis-
covered in Panama.

96 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES
Volume 53, No. 9

MATERIALS AND METHODS

Investigative approaches by the author included a review of herbarium material; field research in
Costa Rica, Panama, Colombia, and Ecuador; anatomical and cytological studies; and observations
made from plants cultivated in greenhouses at the University of Minnesota. Cytological material was
studied from immature flowers fixed in the field with Carnoy’s solution (4:3:1 absolute alcohol, chlo-
roform, glacial acetic acid) and kept as close to 0°C as possible for 1—3 days, after which the material
was stored refrigerated in 70% alcohol. Anthers were teased open, squashed with acetocarmine solu-
tion and permanently mounted in Hoyer’s fluid.

DISTRIBUTION AND HABITAT

The greatest diversity of Monolena species is found in Panama and western Colombia. Five or six
species are found only in the Serrania de Baudo, Rio Atrato basin, Pacific lowlands, and the west flank
of the Andes in Colombia in the departments of Choco, Valle, Cauca and Narino (Uribe 1971). An-
other six species are endemic to Panama and newly described here: Monolena dressleri, M.
grandiloba, M. morleyi, M. multiflora, M. panamensis, and M. trichopoda. In Panama, Monolena oc-
curs along the crest of the cordillera in the vicinity of the continental divide, the Caribbean slope of the
cordillera, and occasionally in the Caribbean lowlands. (Fig. 1)

Monolena primuliflora is the only widespread species of Monolena, occurring from southern
Costa Rica to southern Peru. Monolena guatemalensis is known only from Guatemala, nearly 900 km
from its nearest congener.

Monolena are primarily herbs of wet tropical forests and cloud forests. In Panama they typically
occur in regions of tropical premontane rain forest and tropical wet rain forest (Tosi 1970). In these re-
gions the dry season, from about December through April, is often interrupted by periods of rain; fog
and mist are common throughout the year.

Monolena most often grows as epiphytes on tree trunks, buttresses and branches within three me-
ters of the ground. One collection of M. guatemalensis was reported as growing near the top of a tree.
Most or perhaps all species of Monolena also grow on decaying stumps and logs or occasionally as
terrestrials. Monolena dressleri and M. trichopoda, and some populations of M. grandiloba, M.
multiflora and M. primuliflora, typically grow on rocks in streams.

COMPARATIVE MORPHOLOGY

GROWTH HAsit. — Plants of Monolena are herbaceous; the vegetative shoot axis is condensed
into a fleshy, green to brown rhizome. The rhizomatous habit, anomalous in the Melastomataceae,
creates some difficulty in determining phyllotaxy in the genus. The leaves are basal and are borne in
clusters on the rhizome; each cluster appears to represent a developed bud, with no elongation of its
axis. The first visible structures of the expanding bud are 4-6 small scales that are opposite and
decussate, as would be expected of a Melastomataceae. The foliage leaves then appear, but they are
borne singly, not in pairs, and hence Triana (1871) wrote that they appeared to be alternate. However,
with the leaves appear a few more scales, and sometimes one of these is found approximately opposite
each new leaf, although the opposite position of a given pair changes as the rhizome and leaf base en-
large. Triana proposed that such a paired scale represented a vestigial member of a pair of opposite
leaves.

Roots. — Monolena roots are generally fibrous and are usually entwined in a substrate of vege-
tation and partially decomposed litter. The larger roots are few in number, white, glabrous, and round
in cross-section. The rootlets are more abundant, brown, round to square in cross-section, with a cylin-
drical core and an easily removed outer sheath. The rootlets are glabrous to pubescent. The hairs are

WARNER: MONOLENA IN CENTRAL AMERICA

“euleUR Ul PuajoUOyy JO UONNQIYSIG ‘| TMADIA

rs ars = = = —— al}
ppodoyoia] puajouopy

Sa}awo}ly

a oe ee

002 OOL

pAéopfiynuusd puajouopy
| sisuaupuod vuajouopy
DAOLI] JNU DUajoUuop
DgOjIpupsés DUua;OUOW

1Aaj4OUl DUA]OUOP

eth diac h M

14a]8Sadp DUua]OUo|y

98 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES
Volume 53, No. 9

usually stiff and retrorse and apparently aid in anchoring the plant to the substrate. Rootlets are often
attached to partially decomposed substrate, perhaps affixed by mycorrhizae.

LEAVES. — The leaves of Monolena are well differentiated into petiole and blade. The petioles
are canaliculate and vary from slender in the small, delicate species, to thick and succulent in the more
robust species. The upper blade surface is shiny green, although occasionally the primary veins are
red-purple. In M. dress/eri and its South American allies, the upper blade surface is usually very dark
green with a metallic luster (this group of species includes M. bracteata Gleason, M. cordifolia
Triana, M. dressleri, M. pilosiuscula L. Uribe, and M. pygmeae L. Uribe; it is hereafter referred to as
the Cordifolia complex). In most species the lower blade surface and petiole are red-purple, although
sometimes they are so colored only along the veins, other times in the areoles, and occasionally not at
all. The texture varies from membranous to subcoriaceous. Blade morphology is variable, but gener-
ally consistent within a species. Blades vary from elliptic to lanceolate, ovate, or round. The margins
vary from rarely entire (only in Monolena trichopoda), to more often ciliolate-serrulate and occasion-
ally repand, to crenate-dentate and ciliolate in the Cordifolia complex.

As in most Melastomataceae, foliar venation in Monolena consists of 3—11 longitudinal nerves,
called primaries. Including the midrib, the number of primaries is always odd. The outermost pair of
primaries is usually inconspicuous and converges with the margin in the lower 2/3 of the blade.
Leaves in Monolena vary from nerved to strongly plinerved. Leaves are referred to as “nerved” when
all of the primaries arise from a common point at the base of the blade (see Fig. 2B), or “plinerved”
when one or more pairs arise from the midrib above the base of the blade (see Fig. 3B; Almeda 1978).
Leaves of Monolena species are, with very few exceptions, consistently nerved or plinerved, an easily
observed character useful to distinguish species. The secondary veins sometimes run between and
connect the primaries, or with the tertiary veins form a reticulate pattern. The primary veins, and occa-
sionally the secondary veins, are raised below and sometimes depressed above the leaf surface.

In all but one species of Monolena in Central America the petiole and lower blade surfaces are mi-
nutely puberulent, while the upper surfaces appear glabrous but actually bear scattered minute hairs.
Monolena morleyi is distinct in the genus for being glabrous. Monolena trichopoda exhibits a distinct,
but often caducous fringe of hairs along the ridges of the petiolar canal, and is the only Central Ameri-
can species having large trichomes on the upper blade surface, in addition to the smaller hairs men-
tioned above. However, the large trichomes on the blades of M. trichopoda are rare, apparently found
only on juvenile leaves, and may be present only on some leaves ofa given plant. The South American
species display a greater diversity of trichome types, including some that are peculiar to certain spe-
cies, as in Monolena pilosiuscula, which has sclereids present in the base and shaft of the foliar tri-
chomes.

Sclereids of a different type are found in the leaf blades of some species of Monolena. Monolena
panamensis, M. morleyi, M. grandiloba, M. primuliflora, and some South American species have
sclereids in the hypodermis of the upper surface of the blade. The shape and the abundance of sclereids
in M. primuliflora vary considerably, and they are mostly lacking from plants of lower elevations in
Choco, Colombia, southeastern Peru, and probably in plants of lower elevations in the Talamanca
Mountains of Costa Rica and Panama. Leaves of M. multiflora have sclereids in the hypodermis of
both the upper and lower blade surface and, as in M. primuliflora, the density of sclereids is sharply re-
duced in plants from lower elevations, specifically those populations from Rio Guanche, Colon prov-
ince, Panama. Most plants of M. trichopoda lack foliar sclereids, but where they do occur, they are
found only in the hypodermis of the lower blade surface. Foliar sclereids are completely lacking in M.
guatemalensis and members of the Cordifolia complex.

INFLORESCENCE. — The inflorescence in Monolena is a scorpioid cyme (Triana in Bentham and
Hooker 1867; Cogniaux 1891). More specifically, it is a circinnately coiled cyme, with the flowers al-
ternating from one side to the other, along one side of the pseudoaxis. This type of inflorescence is ap-
parently derived from a dichasium by suppression of successive axes on alternate sides. The peduncle

WARNER: MONOLENA IN CENTRAL AMERICA 99

varies from green to red and from fleshy to very succulent. The number of flowers per inflorescence
ranges from 3 to 35 in the Central American species. Members of the Cordifolia complex in Choco,
Colombia mostly have | or 2 flowers per inflorescence.

Associated with the inflorescence are three types of bracts that can be distinguished by their posi-
tion relative to the flowers: (1) scales, which are borne on the peduncle below the lowermost flower
and its floral bracts; (2) lower floral bracts, which subtend each flower; and (3) lateral floral bracts,
which are borne to the outside of the two rows of flowers. The latter two types of bracts are indistin-
guishable from one another, except by position, and are here collectively referred to as floral bracts.

The scales are somewhat foliaceous, usually green and with venation similar to that of the leaves.
The floral bracts are petaloid, although more succulent than the petals, translucent green to white, and
have parallel venation like that of the petals. The floral bracts are obovate, except in M. multiflora,
where they are broadly ovate to round or reniform. The floral bracts envelop the flower buds until
shortly before the flowers open. In M. multiflora, the floral bracts form a tightly closed sac around the
flower buds.

One lower floral bract subtends each flower, a character consistent throughout the genus. Most
species also bear one lateral floral bract per flower, and four scales per inflorescence. Variation in the
number of scales per inflorescence is seen in the wide-ranging M. primuliflora, which has two or four
(rarely six), and in M. trichopoda which has two or (in some cases) apparently only one. In M.
panamensis, M. multiflora, and M. grandiloba the lateral floral bracts are approximately half the num-
ber of the flowers. These species also bear the greatest number of flowers found in the genus. In M.
guatemalensis, only the two lowermost lateral floral bracts are apparent on the available material.

The floral bracts appear to represent pairs of opposite bracts that have been displaced by the shift-
ing of the flowers to one side of the pseudoaxis and by the swelling and elongation of the pseudoaxis.
Although not readily observable in the floral bracts, such a shift in relative position is evident in the
scales on the peduncle. Initially, the scales are approximately opposite and decussate, but are dis-
placed vertically as the inflorescence elongates.

FLOWERS. — Hypanthium and Perianth. The Monolena hypanthium is obconical to turbinate
and slightly constricted where it surpasses the apex of the ovary; in cross-section, it ranges from round
to triangular. It is 10-nerved and crowned by a highly vascularized region, the torus, which is the point
of insertion of the perianth and stamens.

The calyx lobes are usually connate at the base to form a short tube. The lobes are triangular and
rounded at the apex in most species. The calyx lobes in M. multiflora are rounded at the apex or more
often emarginate, and in M. grandiloba the lobes are exceptionally large, broader at the apex and
deeply obcordate. In some South American species, the calyx lobes are spathulate. The margins of the
calyx lobes are ciliate in all species of Monolena and, in most species, the hypanthium and calyx are
sparsely puberulent. Notable exceptions are M. panamensis and M. morleyi, in which the hypanthium
and calyx lobes are glabrous.

The petals in Monolena flowers are five, convolute in the bud, and free. In most species, the petals
are broadly obovate to oblong and spread to form a shallow cup upon opening. The corolla varies from
pink to white within some species, either as solid colors or white at the base and pink toward the apex.
In other species, it is strictly pink or white. In some plants of M. multiflora, the petals are pink-striped
with translucent white veins. Monolena grandiloba has been described by collectors as having petals
“clear pink” or “bright pink.”

Androecium. The ten stamens in Monolena flowers are disposed in two whorls of 5. The two
whorls are morphologically distinct in that the antesepalous stamens bear a more complex and diverse
connective and are mostly 20-40% larger than the antepetalous stamens. The filaments are white,
dorsiventrally compressed and are declined to one side when the flower opens. The thecae are yellow,
linear, and tipped by a ventrally inclined terminal pore. In bud the stamens are strongly inflexed; the
apex of the thecae rest between the hypanthium and the ovary.

100 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES
Volume 53, No. 9

As in most Melastomataceae, the connective in Monolena is a complex structure, useful in distin-
guishing both the genus and the species. The connective is geniculate just below the thecae, so that the
lower portion of the connective, and the ventral appendage in particular, is positioned between the
thecae and the style. The ventral appendage is the enlarged basal portion of the connective. The dorsal
side of the connective is grooved from below the geniculation to the insertion of the filament. The
groove often continues along the ventral appendage, although it soon becomes indistinct. On the
antesepalous stamens, and occasionally on the antepetalous ones, the upper end of the groove termi-
nates ina small tubercle. The groove embraces the filament when in bud. In the antesepalous stamens
the ventral appendage varies from subglobose in most species to tongue-shaped in M. trichopoda, sad-
dle-shaped in M. dress/eri, and geniculate and pointed in M. grandiloba. The antepetalous stamens
bear only a small linear ventral appendage, or occasionally none at all. In most species of Monolena
the entire connective is yellow. Exceptions are found in M. grandiloba and some populations of M.
multiflora, in which the ventral appendage and dorsal surface on the antesepalous staminal connec-
tives are pink (but turn black on drying).

Gynoecium. The ovary in Monolena is 1/2—3/4 inferior, 3-angled in cross-section, and trilocular.
The apex of the ovary is excavate, with the style emerging from the center. The style is narrow at the
base, swollen near the middle, and slightly contracted and curved below the capitate, papillose stigma.
The numerous ovules are born on axile placentas that are narrowly deltoid in section and intrude into
the locules. The ontogeny and anatomy of the similar gynoecium in Bertolonia marmorata were de-
scribed by Van Heel (1958).

Fruit and seeds. The Monolena fruit is a triquetrous capsule; dehiscing by valves that open at the
top of each locule. The hypanthium is shed as the ovary matures. The seeds are obconic-pyramidate,
with a lateral raphe along most of their length. The surface patterning varies from inconspicuously
roughened to papillate. Seed morphology of Monolena was further described by Whiffin and Tomb
(1972).

FLORAL SYMMETRY AND POLLINATION

Symmetry in flowers of Monolena ts radial with respect to the perianth, but bilateral with respect
to the androecium and gynoecium. The open flowers face at an angle of 0-90 degrees above horizon-
tal. The stamens are declined to the lower side. The style is bent just below the apex so that the stig-
matic surface and thecal pores face toward each other.

As in many taxa with poricidal anther dehiscence, Monolena flowers are probably pollinated by
buzzing insects. In buzz pollination, vibrations created by the insect cause pollen discharge. Although
I observed six species in bloom in the field, few potential pollinators were seen visiting Monolena. In-
sects believed to be bees were observed leaving flowers of Monolena panamensis, and were seen to
inspect, but not land on, flowers of M. pilosiuscula. The insects were presumably foraging for pollen;
no nectar or oil is produced in Monolena flowers, which are also odorless.

An apparent mechanism for self-pollination is present in plants of the Cordifolia complex, in M.
trichopoda, and possibly in other species as well. Late in anthesis the connective folds in sucha way as
to cross the thecae over, or to the side of, the ventral appendage, bringing the thecal pore in contact
with the stigma. The thecae, particularly those of the small stamens, are effectively stuck to the stigma
by the exudate of the latter. The presumed function of this mechanism has yet to be demonstrated ex-
perimentally.

CYTOLOGY

Solt and Wurdack (1980) have reported n = 8 for Monolena dressleri (Dressler 4247), and n = 8
(2n = 16) for plants originating from Cerro Jefe in Panama province (probably Monolena multiflora).

WARNER: MONOLENA IN CENTRAL AMERICA 101

My results are as follows: Monolena dressleri, n=8 (Warner 422); and Monolena multiflora,
n = 8(—9?) (Warner 421).

SYSTEMATIC TREATMENT
Monolena Triana, in Bentham & Hooker, Genera Plantarum. 1(IIT):756 (1867).
NEOTYPE here designated: Monolena primuliflora Hook. f.

Perennial herbs from a fleshy rhizome; root system fibrous; plants mostly puberulent, and occa-
sionally also with long trichomes on the petiole or blade. Scales of the rhizome opposite and
decussate, caducous. Leaves apparently alternate, although opposed by a caducous scale; petiole
canaliculate; blades with 3—11 subparallel primary nerves, the nerves either basal (nerved) or
suprabasal (plinerved), prominent below. Inflorescence a scorpioid cyme or I-flowered scape. Bracts
and scales of the inflorescence mostly caducous, green to white and often translucent, succulent and
foliaceous to more often petaloid; scales mostly subulate and often bearing long apical trichomes; flo-
ral bracts mostly 2 per flower, occasionally only one for some flowers of the inflorescence, ovate to
rarely reniform, entire. Pedicels short. Flowers with a urceolate-campanulate, 10-nerved hypanthium,
crowned by a torus that bears the perianth and stamens. Calyx lobes 5, usually fused at the base to form
a tube, the lobes mostly triangular, rarely spatulate or emarginate or obcordate, the margin ciliate.
Petals 5, free, white to pink, glabrous, obovate to rarely oblong. Stamens 10, glabrous, dimorphic, the
antesepalous whorl distinctly larger than the antepetalous whorl; filaments of both whorls
dorsiventrally compressed and declined to one side of the flower, white; anthers mostly yellow, linear,
2-celled, tipped with a single, ventrally-inclined terminal pore; connective prolonged, geniculate,
usually dorsally tuberculate between the thecae and the insertion of the filament, further extended ba-
sally to form a large appendage on the antesepalous stamens and a small appendage on the
antepetalous stamens. Ovary 1/2—3/4 inferior, glabrous, obconic, trigonous and trilocular, with axile
placentation. Style immersed at base in an ovarial collar, inflated near the middle, slightly contracted
and curved below the capitate, papillate stigma. Ovules numerous, borne on narrow, deltoid placentae
that intrude into the locule. Fruit a triquetrous capsule, dehiscing by 3 apical valves. Seeds
obconic-pyramidate, the surface inconspicuously roughened to papillate.

KEY TO THE CENTRAL AMERICAN SPECIES OF MONOLENA

1. Leaves with all the primary veins diverging from the base of the blade (see Fig. 2B) or with the uppermost pair of pri-
mary veins diverging from the midrib within 1/10 the distance from the base to the apex of the blade; flowers 2-8
(—10) per inflorescence.

2. Pedicels 4-6 x 0.5—0.8 mm; petals 24-33 mm long; petioles mostly drying round or nearly so, and brittle; plants
glabrous except for minute, scant cilia along the margins of leaves and calyx lobes. . . . . . 4. Monolena morleyi
2. Pedicels 2-3 x 0.8—1.2 mm; petals 11-24 mm long; petioles usually drying flat, flaccid or firm, but not brittle;
plants minutely puberulent on underside of the leaves, pedicels and hypanthium.
3. Plants often lithophytic in small streams, occasionally epiphytic on tree trunks and logs or terrestrial; petals
11-15.5 mm long, white; plants to 25 cm; peduncle and axis of the inflorescence in fruit 9-18 cm measured
Tomiie unthestmedicg Cmmemi ea es bc Men ora, es i raf 2a, Aura ueeree 1. Monolena dressleri
3. Plants usually epiphytic on tree trunks and logs or terrestrial, occasionally lithophytic; petals 15—24 mm long,
pink to white suffused with pink or rarely all white; plants to 55 cm; peduncle and axis of the inflorescence in
fruit 19-45 cm measured to the furthest pedicel.
4. Leaves ovate, cordate to rarely obtuse at the base; dried blades membranaceous, dark green above and light
SLEEMDE LOWE GUAtEIMN Alay see ele ett Geko e suet Spey ooh rs eee 3. Monolena guatemalensis
4. Leaves ovate to lanceolate or elliptic, cuneate to obtuse at the base; dried blades subcoriaceous, green to
' brown above, red to green and often dark brown-red along the veins below; Panama and South America.
Ee Para Re dket AE ee AeN econ re tease sh Sole Sat Shae SUE RT Mee are 7. Monolena primuliflora

102 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES
Volume 53, No. 9

1. Leaves with the uppermost pair of primary veins diverging from the midrib 1/5 to 1/3 the distance to the apex of the
blade (see Fig. 3B); flowers 2—35 per inflorescence.
5. Flowers (7—) 12-35 per inflorescence.
6. Leaves ovate to nearly round; pedicels and hypanthium glabrous... .......... 6. Monolena panamensis
6. Leaves elliptic to lanceolate or ovate; pedicels and hypanthium minutely puberulent.
7. Flowers (9—) 14—16 (—26) per inflorescence; calyx lobes (3—) 4-7 mm long, emarginate to broadly obovate;

ventral appendage of antesepalous stamens geniculate and pointed. . . . . . . . 2. Monolena grandiloba
7. Flowers (15—) 20-35 per inflorescence; calyx lobes 1.6—3 mm long, rounded to truncate or emarginate;
ventral appendage of antesepalous stamens subglobose............... 5. Monolena multiflora

5. Flowers 2—8 per inflorescence.
8. Petiole often with two rows of setae 0.9-3.0 mm long along the ridges; leaf blade margin entire, floral bracts

72 immuilons-petalsulsl—liGmnmalll on oem ener ne en 8. Monolena trichopoda
8. Petiole without long setae; leaf blade margin with cilia 0.2 mm long; the floral bracts (10—) 16-20 mm long;
petalsHiS=2Aiminitl One no esate ee eee, he mes eee ee ene ee ay ome 7. Monolena primuliflora

1. Monolena dressleri R. H. Warner, sp. nov.
Figure 2

TYPE: PANAMA, Panama, El Llano-Carti highway, ca. 12—14 km N of El Llano, 350-400 m; 9
January 1973; leaves reddish green above, wine red beneath; R. L. Dressler 4247 (holotype: US!;
isotype: PMA!).

Laminae ovatae, basi cordata usque raro obtusa, nerves vel infirme plinerves; inflorescentia
floribus (2—) 3—6; bracteae florales obovatae; petala | 1—17 mm longa; appendices ventrali selliformes
staminum antesepalum.

Epiphytes on low branches and trunks, and on logs and rocks, or terrestrial. Dried rhizomes 1—7 x
0.6—-1.1 (—2.5) cm. Leaves membranaceous when dry, scantily puberulent throughout, the hairs
0.3—0.5 mm long; petioles red to purple, 2—9 cm long; upper blade surface with a metallic luster, dark
green to reddish green, turning light green on drying; blades 5.5—16 x 3.5—10.5 cm, ovate, cordate to
rarely obtuse at the base, acute to obtuse at the apex, weakly 3—5S (—7) plinerved, the innermost pair of
primary veins diverging from the midrib in an opposite arrangement less than 1/10 of the distance to
the apex; primary veins prominent below; margin ciliolate-serrulate and often dentate-crenate, occa-
sionally repand, the hairs 0.3—0.5 mm long and 20—30/cm. Inflorescence bearing (2—) 3—6 flowers;
peduncle and axis of the inflorescence pink to purple throughout or green toward the apex, together
6.5—13 cm long in flower and 9-18 cm long in fruit when measured to the furthest pedicel, scantily
puberulent, the hairs ca. 0.1 mm long; bracts and scales of the inflorescence caducous, translucent
white to pale green, petaloid, scantily puberulent on both surfaces, the hairs ca. 0.1 mm long; scales 4,
subulate to spatulate to obovate, acute to rounded at the apex, entire to ciliolate-fimbriate with hairs to
1 mm long on the margin; lower and lateral floral bracts each one per flower, all floral bracts 9-11.5
(—15) x 5.0-7.5 mm, obovate, rounded at the apex, entire. Pedicels ca. 2 mm long. Hypanthium ca.
3 mm long, ca. 3.5 (—4.5) mm wide at the torus. Calyx tube absent or to 0.3 mm long; calyx lobes
1.82.0 mm long, 1.8—2.5 mm wide at the base, triangular to oblong, acute to rounded at the apex.
Pedicel, hypanthium, and the calyx scantily puberulent, the hairs ca. 0.1 mm long. Petals white, 1 1-17
x (5—-) 7.5-10.5 mm, obovate, rounded at the apex, entire. Antesepalous stamens: filaments
4.35.0 mm long; thecae yellow, 1.7—2.0 mm long; connective yellow, extended 1.0—1.5 mm be-
tween the thecae and the insertion of the filament, bearing a dorsal tubercle 0.2—0.3 mm long and a
saddle-shaped ventral appendage 0.8—0.9mm long. Antepetalous stamens: filaments (3.2—)
3.6-3.9 mm long; thecae yellow, |.7—2.0 mm long; connective extended 0.6—1.0 mm between the
thecae and the insertion of the filament, the ventral appendage 0.2—0.3 mm long. The connective in
both whorls of stamens bending in late anthesis to bring the thecal pore in contact with the stigma.

103

WARNER: MONOLENA IN CENTRAL AMERICA

imm

| \\ | y)
yy;

VLE.

(==
A AM?
\ i

\ Wy Vitis Poth:

Ani ;
UNAS
(Na

epee

FiGuRE 2. Monolena dressleri R. H. Warner. A. habit; B. leaf blade (abaxial surface); C. inflorescence (fructescence); D. flo-
ral bract; E. flower; F. hypanthium and calyx; G. flower (parts removed); H. antesepalous stamen; I. antepetalous stamen; J.
fruit; K. valve cover from fruit; L. seeds. (A, B, L from Almeda 6509; C, J, K from Antonio 1265; D-I Dressler 4247).

104 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES
Volume 53, No. 9

Ovary ca. 2.0 mm long; style 6.0—6.5 mm long, ca. 1.0 mm wide where expanded. Fruit 1.1—1.2 cm
wide. Seeds ca. 0.7 x 0.6 x 0.5 mm.

DISTRIBUTION AND PHENOLOGY. — This species is known only from the vicinity of the type lo-
cality, from 330 m to 500 m elevation near the border between the province of Panama and the
Comarca of San Blas. Flowering collections have been made in February, March, May and August
and fruiting collections in January February, March, August and December.

PARATYPES. — PANAMA. Panama: El Llano-Carti Rd., 9.6-16 km N of El Llano: Antonio
1265 (CAS), Almeda et al. 6509 (CAS), Busey 380 (MO, US), Croat 33738 (PMA), Folsom & Kauke
1414 (MO), Kennedy et al. 2407 (L, MO), Kennedy & Dressler 2910 (MO), Liesner 1209 (MO), Mori
& Kallunki 6394 (MO), Nee et al. 8761 (MO), Nee et al. 9358 (MO, PMA), Systma & Anderson (MO),
Warner 411, 412, 414, 415, and 422 (all at MIN); San Blas: Nusagandi, El Llano-Carti Rd.: de Nevers
& Gonzalez 3660 (CAS, MO, PMA), van der Werff 7011 (CAS, MO, US).

DISCUSSION. — The Panamanian M. dress/eri is distinguished from all of its South American al-
lies in the Cordifolia complex by its slightly larger size and greater number of flowers; and from indi-
vidual species by the blades being more ovate than round, by the absence of long hairs on the upper
blade surface, and by the calyx lobes being more triangular than spathulate.

ETYMOLOGY. — The holotype of M. dress/eri includes plants in fruit that were collected in the
field and a packet of inflorescences from plants cultivated in Panama by Robert Dressler. Attached to
the sheet is a letter from Dr. Dressler to John Wurdack, dated 27 February 1973, with the following
note, “This plant is quite frequent in the area and grows as epiphyte, lithophyte or terrestrial. The
plants are quite attractive, and so far it seems to be quite easy to cultivate.” Dr. Dressler has collected
nearly every species of Monolena known from Panama, and his are always among the few collections
with well preserved flowers of these difficult to preserve herbs. For these reasons I have named this
species in his honor.

2. Monolena grandiloba, R. H. Warner, sp. nov.

TyPE: PANAMA, Chiriqui, near Cerro Colorado, ca. 10 miles from Chami camp, 8°35’N,
81°45'W, along ridge trail in forest, ca. 1600 m, 15 April 1986, G. McPherson 8938 (holotype: MO!).

Laminae ovatae, basi cuneata usque subcordata, valde plinerves; inflorescentia floribus 14-26;
bracteae florales rotundae usque reniformes; lobi calycis 4-6 (—7) mm longi, emarginati ab obovati
late; petala 26-28 mm longa; appendices ventrali geniculati et acuti staminum antisepalum.

Terrestrial and epiphytic near the base of tree trunks and on rocks in streams. Dried rhizomes 4—9
x 0.5-3.0 cm. Leaves membranaceous, glabrous to scantily glandular puberulent along the secondary
veins on the lower blade surface, the hairs ca. 0.1 mm long; upper blade surface green (on drying may
turn rusty red along the veins near the base of the blade); lower surface green, sparsely in-
dented-punctate. Petioles 3-27 cm long. Blades 10-33 x 4-18 cm, ovate to elliptic, the base cuneate
or truncate to subcordate, the apex acute to caudate, 7—9 (—11) plinerved, the innermost pair of prima-
ries diverging in an opposite arrangment less than 1/4 of the distance to the apex; primary veins promi-
nent below; margin entire and with hairs ca. 0.2 mm long and 5—10/cm. Inflorescence bearing (9—)
14-16 (rarely to 25) flowers; peduncle and axis of the inflorescence together 22-35 cm long when
measured to the farthest pedicel, elongating to 42-48 cm in fruit; peduncle minutely pubescent with
hairs ca. 0.1 mm long; bracts and scales of the inflorescence caducous; scales 2-4; lower floral bracts
one per flower, lateral floral bracts apparently 1/2 the number of lower floral bracts; floral bracts
obovate 1.0—-1.5 x 0.6-1.0 cm. Pedicels 2-4 mm long and 1.2—1.8 mm wide. Hypanthium 5—6 mm
long, 6 mm wide at the torus. Pedicels and hypanthium pubescent. Calyx tube 0.8—1.0 mm long; calyx

WARNER: MONOLENA IN CENTRAL AMERICA 105

lobes (3.0—) 4-7 mm long, (3.0—) 4.0—6.0 mm wide at the broadest, obcordate, the margin minutely
ciliate. Petals white to bright pink, 1.4—2 x 2 cm, obovate, rounded at the apex, entire. Antesepalous
stamens: filament 4.5—-5.0 mm long; thecae yellow, 3.5—3.8 mm long; connective yellow with the
ventral appendage and dorsal surface black on dried material up to and including the tubercle; connec-
tive extended 2.0—2.4 mm between the thecae and the insertion of the filament, bearing a dorsal tuber-
cle ca. 0.2-0.4 mm long, and a ventral appendage 1.0—1.2 mm long, geniculate and pointed.
Antepetalous stamens: filaments 3.84.6 mm long; thecae yellow, 3.0 mm long; connective yellow,
extended 1.5 mm between the thecae and the insertion of the filament, the ventral appendage
0.5—0.6 mm long. Ovary 3.0—3.2 mm long; style 7—9 mm long, 1.0 mm wide where expanded. Fruits
(immature) 1.3 cm wide. Seeds not available.

DISTRIBUTION AND PHENOLOGY. — Monolena grandiloba is endemic to Panama, where it is
found in the vicinity of the type collection at an altitude of 1500-1600 m along the continental divide
between Chiriqui and Bocas del Toro and Veraguas from 400—1350 m in forests and along streams
crossing the road between Alto Piedra and Calovébora and up to the ridges of Cerro Tute. This species
has been collected in flower in March, April, May and June and in fruit in March, April and June.

PARATYPES. — PANAMA. Chiriqui: Cerro Colorado Mine, near higher elevation camp,
1,500 m, Antonio 4867 (MO); along continental divide road 13—15 km beyond intersection in Hato
Chami, Gomez & Warner 511 (PMA); Veraguas: Cerro Tute, ca. 10km NW of Santa Fe,
800-1350 m. Antonio 3950 (MO), Folsom & Edwards 3362 (CAS), Knapp & Dressler 5424 (CAS,
MO), Mori 6741 (MO); in forest and along streams crossing road between Alto Piedra and
Calovébora, 400—900 m, Croat 27458 (MO, PMA), Dressler 4998 (US 2 sheets), Nee 11236 (MO).

DISCUSSION. — Monolena grandiloba is distinguished from the other species of Monolena by its
floral bracts ovate and not enclosing flower in a sac; flowers 5—15 (rarely to 25); calyx lobes broadly
obovate, typically 4-7 mm long and 4.0—6.0 mm at the widest; and stamen appendage pointed, some-
times geniculate and often black on dried material.

This species is apparently related to Monolena multiflora, based on the large number of flowers
relative to other species of Monolena, the pubescence on the hypanthium and calyx lobes, and the
emarginate to obcordate calyx lobes.

ETYMOLOGY. — The specific epithet refers to the lobes at the end of each calyx segment. On
some specimens the calyx segments are so broadly obcordate that pressed on herbarium sheets the
lobes look like a pair of mouse ears.

3. Monolena guatemalensis Donn. Sm. Bot. Gaz. 42:294. 1906.

TYPE: GUATEMALA, Alta Verapaz, trail from Senju to Actala, moist bank along trail; succu-
lent, flowers pink; 17 January 1905. William R. Maxon & R. Hay 3331 (Lectotype designated here:
US!).

Epiphytes high in trees, on stumps, or terrestrial. Dried rhizomes (4—) 6.5—15 x 0.7—2.0(-3.0) cm.
Petioles (2.5—) 4-24 cm long; fresh blades “fleshly-membranous, rich green with prominent
areolations above, purplish-orchis or suffused with pale silvery-green beneath” (Steyermark 41922);
dried blades membranous, light green below and dark green above, (6.5—) 8-33 x (2.4—) 5-21 cm,
ovate, cordate at the base, acute to subacuminate at the apex, with entire or sometimes crenate mar-
gins, weakly 5—7 (9) plinerved, the innermost pair of primary veins diverging from the midrib in an
opposite arrangement in the lower 1/10 of the blade, the primary veins prominent below; leaves
puberulent throughout with hairs 0.2—0.3 mm long, the margin with hairs 0.3—0.5 mm long and ca.
4-20/cm. Inflorescence bearing 3—7 flowers; peduncle and axis of the inflorescence together
12-20 cm long in flower and 19—33 cm long in fruit when measured to the furthest pedicel, scantily
puberulent with hairs ca. 0.2 mm long; bracts and scales of the inflorescence caducous, petaloid, en-

106 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES
Volume 53, No. 9

tire, scantily puberulent on both sides with hairs ca. 0.1 mm long; scales 4, 7-12 x (2—) 3-5 mm,
subulate to obovate and rounded at the apex, occasionally with a few hairs near the apex; lower floral
bracts one per flower; lateral floral bracts 2, these approximately opposite the 2 lowermost flowers; all
floral bracts (6—) 10-15 x (3—) 7-10 mm, obovate, rounded at the apex. Pedicels 2—3 mm long.
Hypanthium ca. 3.5 mm long, 3.5—4.5 mm wide at the torus. Calyx tube 0.1—0.2 (-0.5) mm long; ca-
lyx lobes 1.5—2.3 mm long, 2.0—2.2 mm wide at the base, triangular, rounded at the apex. Pedicel,
hypanthium, and margin and both sides of the calyx lobes scantily puberulent with hairs ca. 0.1 mm
long. Petals pink, 22-24 x 13-20 mm, obovate, rounded at the apex, entire. Antesepalous stamens: fil-
aments 4.0-4.5 mm long; thecae ca. 3 mm long; connective extended |.0—1.3 mm between the thecae
and the insertion of the filament, bearing a dorsal tubercle ca. 0.2 mm long and a subglobose ventral
appendage 0.7—0.9 mm long. Antepetalous stamens: filament 3.24.0 mm long; thecae ca. 2.7 mm
long; connective extended ca. 1 mm long between the thecae and the insertion of the filament, the ven-
tral appendage 0.2-0.3 mm long. Ovary ca. 2.5 mm long; style 6.0—6.5 mm long, 0.3—0.6 mm wide
where expanded. Fruit ca. 1.4 mm wide. Seeds ca. 1.1 x 0.6 x 0.4 mm.

DISTRIBUTION AND PHENOLOGY. — This species is endemic to Guatemala and is disjunct by ca.
900 km from its nearest congener in Costa Rica, so far as is now known. It occurs in central Guatemala
at (300—) 900-1500 m elevation. Monolena guatemalensis has been collected in flower in January,
April and May, in fruit in January and March.

ADDITIONAL SPECIMENS EXAMINED. — GUATEMALA. Alta Verapaz: Secanquim, trail to
Secoyocta, Goll 158 (US); vicinity of Sepacuite, Cooks & Doyle 32 (US); near the finca Sepacuite, Se
Shun, Cook & Griggs 106 (Syntype: US); near finca Sepacuite, Cook & Griggs 515 (Syntype: US);
Between Sepacuite and Sicanquim, Pittier 3/4 (Syntypes: US & NY, photos at GH & MO); Quiche:
Cerro Putul, “Zona Reyna,” Skutch 1826 (F, GH, NY, US); Izabal: Cerro San Gil, Stevermark 41922
(EF. NY OUS 2 sheets):

DISCUSSION. — The lectotype designated here was selected because it has the best preserved
flowers of the specimens cited by Donnell Smith in the protologue. However, the petioles are some-
what longer than typical.

Monolena guatemalensis is distinguished by a combination of characters not seen elsewhere in
the genus. The blades are ovate, nerved to slightly plinerved, and lacking sclereids as in members of
the Cordifolia complex. However, these plants differ from those of the Cordifolia complex by their
larger size, fewer lateral floral bracts relative to the number of flowers, and antesepalous stamens with
a subglobose ventral appendage. The shape of the staminal appendages and the low number of floral
bracts suggest a relationship to M. panamensis. The overall pubescence of M. guatemalensis is of
slightly longer hairs than typical for the genus.

4. Monolena morleyi R. H. Warner, sp. nov.

Type: PANAMA, Coclé, 7 km north of El Copé, Forgotten Hill, area surrounding Rivera Saw-
mill; 650-850 m; 5 Nov 1977; epiphytic herb, petals pink, stamens yellow; J. P. Folsom 6204
(holotype: US!; isotypes: CAS! and MO!).

Plantae glabrae; laminae ovatae usque fere rotundae, basi cordata, nerves vel infirme plinerves;
inflorescentia floribus 4—8 (—10); bracteae florales obovatae; pedicelli florum 4—6 mm tenues; petala
24-33 mm longa; appendices ventrali subglobosi staminum antisepalum.

Epiphytic herbs. Dried rhizomes 4-10 cm x 1-2 cm. Leaves glabrous; petioles nearly round and
brittle when dried, deeply caniculate, red, turning red-brown on drying, 4-13 cm long; blades 7-20 x
6-17 cm, ovate, cordate and sometimes oblique, subcaudate to acute at the apex, subcoriaceous, 7—9

WARNER: MONOLENA IN CENTRAL AMERICA 107

(—11) -nerved, to slightly plinerved, the innermost pair of primary veins diverging from the midrib in
an opposite arrangement up to 1/10 of the distance to the apex; primary veins very prominent below,
raised into prominent ridges near the base of the blade; margin faintly ciliolate-serrulate, the hairs
0.5—1.5 mm long and 5—8/cm; on drying the upper blade surface dark red-tan to green, the lower sur-
face indented-punctate, tannish green to brown and with the veins red-brown. Inflorescence with 4-8
(—10) flowers; peduncle pink, wiry when dried, peduncle and axis of the inflorescence together
15—22 cm long in flower and 32 cm in fruit when measured to the furthest pedicel, glabrous; scales
(2—) 4, white; lower floral bracts one per flower, the lateral floral bracts approximately halfas many as
the lower floral bracts, all floral bracts 1.7—2.2 x 1.1—1.8 mm, obovate, rounded to emarginate at the
apex, entire. Pedicels 4-6 mm long, 0.5—0.8 mm wide at the narrowest point. Hypanthium 3-4 mm
long, 4.0-4.5 mm wide at the torus. Calyx white; calyx tube 0.4—0.8 mm long; calyx lobes
2.5—2.7 mm long, 2.2—2.5 mm wide at the base, triangular and rounded at the apex to oblong with the
apex truncate to rarely emarginate; margin of the calyx lobes scantily ciliolate with hairs ca. 0.1 mm
long, the pedicels, hypanthium and calyx otherwise glabrous. Petals white, pink or lavender, 24—33 x
24 mm, obovate, rounded at the apex, entire. Antesepalous stamens: filaments 4.0—-4.8 mm long;
thecae yellow, 2.4—3.0 mm long; connective extended |.2—2.0 mm between the thecae and the inser-
tion of the filament, bearing a dorsal tubercle 0.2 mm long and a subglobose ventral appendage
1.0—-1.2 mm long. Antepetalous stamens: filaments 3.8—4.0 mm long; thecae yellow, 2.0—2.5 mm
long; connective extended |.0—1.2 mm between the thecae and the insertion of the filament, the ven-
tral appendage 0.3—0.6 mm long. Ovary ca. 2.0 mm long; style 5.0—7.0 mm long and ca. 0.8 mm wide
where expanded. Fruits 1.3—1.4 mm wide. Seeds ca. 0.8 x 0.8 x 0.5 mm.

DISTRIBUTION AND PHENOLOGY. — Monolena morleyi is endemic to Panama, where it is known
only from the vicinity of El Copé in the province of Cocle. It grows in cloud forest from 700-1300 m
elevation, including the wind-swept ridge along the continental divide. Flowering material has been
collected in September, October, November, February and April. Fruiting plants have been collected
in January.

PARATYPES. — PANAMA. Cocleé: continental divide above El Copé, region of Alto Calvario
and El Potroso (Rivera) sawmill, Croat 44681 (MO), Croat 67561 (MO), Folsom & Collins 1537
(MO), Folsom & Collins 6535 (MO), Hammel 2427 (MO), McPherson 7673 (MO), Sytsma 1815
(MO).

DISCUSSION. — This species of Monolena is unique in the genus for being entirely glabrous; peti-
oles brittle on dried material; pedicels longer and more delicate than other species of Monolena, and
having the largest petals in the genus. Monolena morleyi is most closely related to M. panamensis.
Both species have ovate to cordate leaves; glabrous pedicels and hypanthium; long pedicels relative to
other species of Monolena; large floral bracts that only partially envelop the flower buds; and large,
showy flowers. The two species are distinguished by the unique characters of M. morleyi mentioned
above and the larger number of flowers per inflorescence of M. panamensis.

While leaves of this species were not cleared, based on the texture, color and surface features of
the leaf blade are similar to M. multiflora; thus I suspect that foliar sclereids are abundant and associ-
ated with the lower hypodermis, and possibly with the upper hypodermis as well.

ETYMOLOGY. — This species is named for the late Dr. Thomas Morley, an expert on the genera
Mouriri and Votomita (Melastomataceae), dedicated conservationist, and teacher of plant taxonomy
to generations of students at the University of Minnesota.

108 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES
Volume 53, No. 9

5. Monolena multiflora, R. H. Warner, sp. nov.
Figure 3

TYPE: PANAMA, Panama, La Envida, region of Cerro Jefe; flowers pale pink; 18 April 1971;
R. L. Dressler & N. H. Williams 3950 (holotype: US 2849129!; isotypes: MO!, PMA!, US 2849130!).

Laminae ellipticae usque ovatae, basi cuneata usque obtusa, valde plinerves; inflorescentia
floribus (15—) 20-35; bracteae florales rotundae usque reniformes; petala (10—) 15-29 mm longa; ap-
pendices ventrali subglobosi staminum antisepalum.

Epiphytes on tree trunks and low branches, and on stumps and rocks. Dried rhizomes 3.0—8.0 x
0.6—2.0 cm. Plants mostly glandular puberulent, the hairs most conspicuous on the margin of the leaf
blade and along the veins on the lower blade surface; hairs 0.3—0.5 mm long and mostly ca. 30/cm
along the blade margin, elsewhere 0.1—0.2 (—0.3) mm long; petals, androecium, gynoecium and fruit
glabrous. Leaves subcoriaceous; upper blade surface green, often turning red-brown on drying; lower
surface rarely green throughout, mostly red, particularly along the veins, turning green with dark
brown to red veins and often becoming indented-punctate on drying. Petioles red, 4-20 cm long;
blades 14-30 x 3.5—18 (—23) cm, elliptic to ovate-lanceolate, cuneate to obtuse and often decurrent at
base, acute to acuminate at apex, strongly 5—7 (9) plinerved, the innermost pair of primaries diverg-
ing in an alternate to rarely opposite arrangement from (rarely 1/10) 1/5 to 1/3 of the distance to the
apex; primary and occasionally secondary veins sunken above and prominent below; margin
ciliolate-serrulate and sometimes repand. Inflorescence bearing (15—) 20-35 flowers; peduncle and

axis of the inflorescence red and succulent, together 10.5—43 cm long when measured to the farthest
pedicel, often bearing both flowers and fruit; bracts and scales of the inflorescence caducous, translu-
cent pink to white or green; scales 4 (-6), 8-12 x 4-8 mm, obovate, rounded at the apex, the margin
entire or occasionally ciliate near the apex, the hairs ca. 0.3 mm long; lower floral bracts one per
flower, lateral floral bracts approximately half as many as the lower floral bracts, all floral bracts
9.5-16 x 9.0-16 mm, broadly obovate to round or reniform, rounded to emarginate at the apex,
strongly concavo-convex, entire; the floral bracts form a pouch that envelops the flower buds until
shortly before the flowers open. Pedicels 0.5—2.0 mm long, 0.8—1.5 mm wide. Hypanthium green or
pink, 2.5-4.2 mm long, 3.0—4.0 (—5.0) mm wide at the torus. Calyx tube 0.3—0.6 mm long; calyx lobes
green to pink, 1.6—2.5 (—3.0) mm long, 2.0—2.5 mm wide at the base, ranging from triangular with a
central ridge that terminates just below the apex, to truncate, emarginate or slightly obovate. Petioles,
hypanthium and calyx scantily puberulent. Petals white to cream or pink and often with translucent
veins, (10—) 15—20 x (6—) 8-11 mm, obovate to oblong, rounded at the apex, entire. Antesepalous sta-
mens: filament 3.44.5 (—5.0) mm long; thecae yellow, 2.3—3.0 mm long; connective entirely yellow,
or (in Colén and Coclé) bicolored with the ventral appendage and dorsal surface pink up to and includ-
ing the tubercle; connective extended 1.2—1.5 (—2.1) mm between the thecae and the insertion of the
filament, bearing a dorsal tubercle ca. 0.1—0.2 mm long, and a subglobose ventral appendage 0.6—0.8
(—1.3) mm long. Antepetalous stamens: filaments 3.1—3.6 mm long; thecae yellow, 2.0—2.4 mm long;
connective yellow, extended 0.8—1.0 mm between the thecae and the insertion of the filament, the
ventral appendage 0.2—0.3(—0.5) mm long. Ovary 3.0—3.2 mm long; style (5.0—) 5.5—6.5 mm long,
0.7—-1.7 mm wide where expanded. Fruits 1.2—1.5 cm wide. Seeds ca. 0.8 x 0.5 x 0.4 mm.
DISTRIBUTION AND PHENOLOGY. — Monolena multiflora is endemic to Panama. East of the Pan-
ama Canal it occurs in Panama province from 1000 m in the Cerro Jefe region east to the area where
the provinces of Panama and Colon border San Blas at elevations of 400-500 m and in Colon from
600 min the Santa Rita Ridge area to near sea level along the Rio Guanche. West of the Panama Canal,

109

WARNER: MONOLENA IN CENTRAL AMERICA

itt

| mtn .
Ni =

eo cy
LY ii,
Y, > Y Y Uy if Uh MS

Yat an

); C. inflorescence (primarily with

(parts removed); H. antesepalous stamen; I. antepetalous

FiGurE 3. Monolena multiflora R. H. Warner. A. habit; B. leaf blade (abaxial surface

fruits); D. floral bract; E. flower; F. hypanthium and calyx; G. flower
stamen; J. fruit; K. valve cover from fruit; L. seeds. (A, B from Dressler 4132: C, J-L from Sytsma et al. 2471.; D-I from

Dressler 4251.)

110 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES
Volume 53, No. 9

M. multiflora occurs in Coclé from 800 m near Cope to less than 100 m near Coclé Del Norte in Colon.
This species has been collected in fruit and flower every month of the year.

PARATYPES.— PANAMA. Coclé: continental divide between Llano Grande and Cascajal,
Knapp 1966 (CAS, PMA), Sytsma et al. 4422 (MO), 4393 (CAS), Sytsma 3846 (CAS), Sytsma 3903
(CAS); continental divide N of Penonomée, road to Coclesito, Antonio 3068 (CAS), Croat 49239
(CAS), D’Arcy & Hammel 12284 (CAS, MO), Hammel 1630 (CAS, MO), Hammel 7211 (MIN),
Churchill et al. 4155 (CAS, PMA); between Rio Blanco and Cano Sucio, Davidse & Hamilaton
23623 (PMA), Sytsma et al. 2471 (CAS), Antonio 3674 (MO); trail from Cano Sucio to waterfall of
Rio Tife, Knapp 3767 (MO); Atlantic side of continental divide above El Cope, Knapp & Dressler
3462 (CAS); Rio San Juan above confluence with Rio Tife, Hammel 3445 (CAS); Colon: Coclé del
Norte, east of town, Hammel 4514 (MO); 6 km S of Porto Bello, along Rio Guanche, 5—20 m, Nee &
Gentry 8707 (MO), Kennedy & Foster 2176 (F, MO, NY, US), Foster 2778 (F, MIN, PMA), Warner
427 (MIN), Maas et al. 1576 (F, MO, US), Dressler 4140 (US), Dressler 4242 (US), Dressler 4301
(PMA, US); Rio Piedras, Santa Rita Ridge, Antonio 3770 (cultivated material), Antonio 3771 (MO),
Hammel 6353 (MO); Rio Gatun, Antonio 3827 (MO); south approach to Cerro Bruja from Rio
Escandaloso, Antonio 1340 (MO), Hammel 3109 (CAS), Hammel 3187 (CAS); Panama: 6 miles
above Goofy Lake on road to Cerro Jefe, Croat 15214 (MO); Cerro Jefe, Correa & Dressler 558
(PMA), Almeda & Nakai 3462 (CAS), Almeda et al. 5832 (CAS), Croat 67071 (CAS, PMA),
Dressler 5138 (US), Henslow & McPherson 1003 (PMA), Mori 6519 (MO), Sytsma et al., 2852
(CAS); La Eneida, region of Cerro Jefe, 650 m, Dressler 4132 (PMA, US), 4251 (US); woods around
La Eneida, Luteyn & Kennedy 1768 (F, MO); beyond La Eneida, Correa & Dressler 804 (GH, MO,
NY, PMA, US); Altos del Rio Pacora, Dressler 5287 (PMA), Lewis et al. 2265 (GH, MO); Campo
Tres, 3 mi NE of Altos de Pacora, Liesner 547 (CAS, MO); 5—10 km NE of Altos de Pacora, on ridge
top, ca. 600 m, Mori & Kallunki 3422 (MO); San Blas: continental divide 5-10 km west of El
Llano-Carti Road, de Nevers & Herrera 4495 (CAS, MO); Cerro Brewster, de Nevers et al. 4032
(CAS, MO); Yar Bired, continental divide between Cangandi and San José, de Nevers & Herrera
7001 (MO, CAS).

DISCUSSION. — Monolena multiflora is distinguished by the leaves ovate to lanceolate or elliptic
and strongly plinerved, the large number of flowers (15—35) per inflorescence, and the floral bracts
that envelop the buds in a pouch.

There are three distinct populations of M. multiflora. Plants from the population east of the Pan-
ama Canal in the region of Cerro Jefe east into San Blas are usually distinguishable by leaves turning
iron red on drying, presumably related to the abundance of sclereids; plants from this region have the
highest density of foliar sclereids of all the specimens examined to date.

In Colon, also east of the canal, M. multiflora most often occur as lithophytes along streams and
tend to be of smaller stature and with leaf blades narrow and delicate (presumably reflecting the very
low density of foliar sclereids recorded from plants in this region). The lithophytic and riverine habi-
tat, smaller stature and more delicate leaves of M. multiflora in the Santa Rita Ridge area are character-
istics reminiscent of M. trichopoda. However, these two species can be distinguished by the larger
number of flowers and the ciliolate-serrulate margin of the leaf blades on M. multiflora, and by the two
rows of trichomes on the petioles of M. trichopoda. Specimens from Rio Guanche suggest that plants
in this particular area rarely set fruit.

West of the Panama Canal, M. multiflora is distinguished by strongly decurrent leaf bases and
flower buds enclosed in a particularly pronounced pouch formed by the floral bracts. Monolena
multiflora 1s sympatric with M. trichopoda in the province of Coclé (along the Rio Blanco), although
their habitats are distinct. Monolena trichopoda grows primarily on rocks and logs in and along fast
running rivers, but is seldom found in the adjacent forest where M. multiflora grows 1n this region.

ETYMOLOGY. — The specific epithet acknowledges that this species has the most flowers per in-
florescence of all Monolena species.

WARNER: MONOLENA IN CENTRAL AMERICA 11]

6. Monolena panamensis R. H. Warner, sp. nov.

TYPE: PANAMA, Coclé, La Mesa, 4 km N of El Valle, 875 m; terrestrial, petals pink; 12 Febru-
ary 1974; M. Nee & Hale 9638 (holotype: US!; isotype: MO!).

Laminae ovatae usque fere rotundae, basi cordata vel truncata vel raro obtusa et interdum
obliqua, modice plinerves; inflorescentia floribus 12-20; bractea florales obovatae; petala
(12—)14—24 mm longa; appendices ventrali subglobosi staminum antisepalum.

Epiphytes on tree trunks, or on stumps and logs, or terrestrial. Dried rhizomes 4—9 x 0.6—2.0 cm.
Leaves subcoriaceous, rarely glabrous, usually scantily puberulent throughout and most conspicu-
ously so along the veins of the lower surface, the hairs 0.1—0.2 mm long; petioles fleshy, caniculate,
red, 6-29 cm long; upper blade surface green and occasionally with the margin red; lower surface
light green when fresh, indented-punctate and usually turning tan or rarely pinkish on drying; blades
7.5—23 x 8-23 cm, ovate to nearly round, cordate and sometimes oblique, truncate or rarely obtuse at
the base, acuminate to rarely subcaudate at the apex, weakly 5—7 (—9) plinerved, the innermost pair of
primary veins diverging from the midrib in an opposite arrangement up to 1/5 of the distance to the
apex, or (in plants from Cerro Trinidad) more prominently 9 (—11) plinerved, the innermost pair of pri-
mary veins diverging from the midrib in an opposite arrangement up to 1/4 of the distance to the apex,
primary veins prominent below; margin apparently entire but actually faintly ciliolate-serrulate, the
hairs 0.3—0.5 mm long and 2-4/cm. Inflorescence bearing 12—20 flowers; peduncle and axis of the in-
florescence red to purple, succulent, together 15—30 cm long in flower and (15—) 30-42 cm long in
fruit when measured to the furthest pedicel, very scantily puberulent with hairs ca. 0.1 mm long;
bracts and scales of the inflorescence caducous, translucent green to white, petaloid, very scantily
puberulent on the adaxial surface with hairs ca. 0.1 mm long; scales 4, ca. 2.1 x ca. 0.9 mm, obovate,
round at the apex, the margin entire and occasionally with a few hairs near the apex, the hairs ca.
0.3 mm long; lower floral bracts one per flower, the lateral floral bracts approximately half as many as
the lower floral bracts, all floral bracts 1.4—-1.5 x 1.0—-1.1 mm, obovate, rounded at the apex,
concavo-convex, entire. Pedicels 2—3.5 mm long, 0.8—1.2 mm wide at the narrowest point.
Hypanthium 3—4 mm long, 4.0-4.5 mm wide at the torus. Calyx tube 0.4—0.6 mm long; calyx lobes
2.52.7 mm long, 2.2—2.5 mm wide at the base, triangular and rounded at the apex to oblong with the
apex truncate to rarely emarginate. Margin of the calyx lobes scantily ciliolate with hairs ca. 0.1 mm
long, the pedicels, hypanthium and calyx otherwise glabrous. Petals pink to lavender, (12—) 14-24 x
8-10 mm, obovate, rounded at the apex, entire. Antesepalous stamens: filaments 2.8—6 mm long;
thecae yellow, 2.24 mm long; connective yellow, extended 1.6—2.5 mm between the thecae and the
insertion of the filament, bearing a dorsal tubercle 0.2—0.3 mm long, and a subglobose ventral append-
age 0.8-1.5 mm long. Antepetalous stamens: filaments 2.6—3.2 (4.0) mm long; thecae yellow,
2.0—3.0 mm long; connective yellow, extended 1.0—1.4 mm between the thecae and the insertion of
the filament, the ventral appendage 0-4—0.5 mm long. Ovary ca. 2.4 mm long; style 5.5—6.0 mm long,
and ca. 1.0 mm wide where expanded. Fruits 1.1—1.2 mm wide. Seeds ca. 0.8 x 0.8 x 0.5 mm.

DISTRIBUTION AND PHENOLOGY. — Monolena panamensis 1s endemic to Panama. Typical
plants are found in the mountains north of El Valle de Anton, Coclé, from 800-1100 m elevation. A
distinct sort of Monolena panamensis is known only from Cerro Trinidad, province of Panama, at
950 m elevation. Flowering collections have been made in every month except August and October,
fruiting collections from January through September.

PARATYPES. — PANAMA. Cocle: along road to and in forest at La Mesa, 2.8—3.4 miles NW of
El Valle de Anton, Correa et al. 4245 (PMA), Luteyn 4069 (F); La Mesa, Warner 404 through Warner
410 (MIN); La Mesa, ca. 2 km W of Cerro Pilon, Croat 37432 (MO), de Nevers et al. 6339 (CAS,
MO); Sullivan 505 (CAS, MO); Cerro Pilon, ca. 5 km NE of El Valle, Croat 14346 (F, MO NY),

ily PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES
Volume 53, No. 9

Dwyer et al. 4481 (MO), Dwyer & Lallathin 8597 (NY), Kirkbride 1072 (MO-2 sheets, NY), Liesner
770 (MO), Mori et al. 6570 (MO, US); hills N of El Valle de Anton, Dressler 4081 (US-2 sheets),
Hamilton et al. 4124 (MO), Lewis et al. 1773 (MO); Cerro Gaital Caracoral, Croat 67256 (CAS, MO,
PMA, US), Duke & Dwyer 15137 (MO), Dwyer & Correa 8840 (MO), Knapp 1063 (CAS, MO, US),
Knapp 5309 (CAS), Knapp & Dressler 4899 (CAS), McPherson 11218 (MO), Miller et al. 783 (CAS,
PMA, US), Sytsma 3744 (MO), Sytsma 3772 (MO), Sytsma 4070 (CAS); Cerro Pajita, region N of El
Valle de Anton, Allen 4494 (MO); Panama: trail to top of Cerro Pelado, 1,000 m elevation, Antonio
1081 (CAS, MO); Capira, NE side or Cerro Trinidad, 950 m elev., Foster 2118 (F, MIN, MO, PMA,
US).

DISCUSSION. — The distinguishing characters of M. panamensis include ovate leaves that are
weakly plinerved, the inflorescence with 12—20 flowers, lateral floral bracts half as many as the lower
floral bracts, obovate floral bracts, and the glabrous hypanthium. Foliar sclereids are associated only
with the lower hypodermis and are moderately abundant in typical plants, but are rare in plants from
Cerro Trinidad.

The plants from Cerro Trinidad differ from typical plants in a number of points. In particular, the
petioles are mostly longer (to 29 cm) than typical plants (to 16 cm); the leaf blades are rounder and
more strongly cordate at the base, have 9 primary veins (compared to 5—7 primary veins in typical
plants), and the blades are more prominently plinerved; and the stamens are about 50% larger than in
typical plants.

ETYMOLOGY. — The specific epithet acknowledges that this species is endemic to Panama.

7. Monolena primuliflora Hook. f., Bot. Mag. 1870:5818 (February 1, 1870).
LECTOTYPE here designated: Plate 5818 in the Botanical Magazine.

Bertolonia primuliflora H. H. Dombrain, Floral Mag. 9:471—472 (1870). Lectotype here desig-
nated: Plates 471—472 in the Floral Magazine. The Floral Magazine cannot be specifically dated, but
must have been published at about the same time as the Hooker description. In his treatment of
Monolena primuliflora Hooker noted that the synonym Bertolonia primuliflora was commonly used
among horticulturalists, but he did not mention Dombrain’s article.

Epiphytic or terrestrial. Dried rhizomes 3—9 x 0.6—3 cm. Plants mostly glandular-puberulent, the
hairs most conspicuous along the blade margin and along the veins on the lower blade surface; hairs
0.1—0.2 mm long, ca. 20/cm along the margin; juvenile leaves occasionally bearing hairs 2.0-2.5 mm
long on the adaxial surface. Leaves mostly subcoriaceous, plants from Cerro Matama in Costa Rica
are membranous; petioles red to green, 7-20 cm long; upper blade surface green, often turning brown
on drying; lower surface red to green often turning dark brown-red along the veins on drying; blades
10-25 cm long, 4-17 cm wide, ovate, lanceolate to nearly elliptical, cuneate to obtuse at the base,
acuminate at the apex; blades (3—) 5—7 (—9)-nerved or plinerved, the innermost pair of primary veins
diverging from the midrib in an opposite arrangement within 1/10 of the distance to the apex (on
plants from eastern Panama and South America), or within 1/5 to 1/3 of the distance to the apex (on
plants from the Talamanca Mountains of western Panama and adjacent Costa Rica); primary veins
prominent below; margin ciliolate-serrulate; juvenile leaves occasionally bearing hairs 2.0—2.5 mm
long on the adaxial surface. Inflorescence bearing 2—6 (—7) flowers; peduncle and axis of the inflores-
cence red, succulent, together 6-19 cm long in flower and 27-45 cm long in fruit when measured to
the furthest pedicel; scales of the inflorescence 2 or 4, 7-12 x 1.5—5.0 mm, subulate to elliptic to
obovate, acute to rounded at the apex, entire and occasionally with a few marginal hairs 0.3—0.8 mm
long; lower and lateral floral bracts each one per flower, all floral bracts 10-20 mm long (10-16 mm
long on plants from eastern Panama, 16-20 mm long on plants from western Panama and Costa Rica),

WARNER: MONOLENA IN CENTRAL AMERICA 113

(5—) 9-14 mm wide, obovate, rounded at the apex, concavo-convex, entire. Pedicels 2~3 mm long.
Hypanthium 2.5—3.5 (—5.0) mm long, 34.5 mm wide at the torus. Calyx tube 1.5—4.0 mm long, calyx
lobes 1.5—3.0 mm long, 2.5—3.2 mm wide at the base, triangular to oblong, rounded to emarginate at
the apex. The pedicel, hypanthium and calyx scantily puberulent, the hairs ca. 0.1 mm long. Petals
white to pink or purple, 15—24 x 12-15 mm, obovate, rounded at the apex, entire. Antesepalous sta-
mens: filaments 3.5—5.0 mm long; thecae yellow, 2.0-2.5 mm long; connective yellow, extended
1.0-1.5 mm between the thecae and the insertion of the filament, bearing a dorsal tubercle
0.1—0.2 mm long and a subglobose ventral appendage 0.8—1.2 mm long. Antepetalous stamens: fila-
ments 2.84.2 mm long; thecae yellow, 1.6—1.9 (—2.3) mm long; connective yellow, extended
0.6—1.0 mm between the thecae and the insertion of the filament, bearing a dorsal tubercle ca. 0.1 mm
long and a subglobose ventral appendage 0.3—0.5 mm long. Ovary ca. 2.4 mm long; style (3.5—)
6.0—7.0 (7.8) mm long, ca. 1.8 mm wide where expanded. Fruit 1.5—1.7 cm wide. Seeds ca. 0.9 x 0.5
x 0.4 mm.

DISTRIBUTION AND PHENOLOGY. — This is the most wide-ranging species of Monolena. It oc-
curs from eastern Costa Rica to southern Peru and adjacent Brazil (Acre), at elevations ranging be-
tween 300-1900 meters. In Central America M. primuliflora has been collected in flower and fruit
from January through August.

ADDITIONAL SPECIMENS EXAMINED. — COSTA RICA. Limon: Asuncién, Rio Segundo,
300-600 m, Gomez et al. 23426 (CAS, MO, US), Gomez & Herrera 23472 (CAS, US). PANAMA.
Bocas del Toro: continental divide trail near Fortuna Dam, 1200-1300 m, A/meda et al. 6257 (CAS);
Continental Divide above Quebrada Arena, Carretera del Oleoducto, IRHE Fortuna Hydroelectric
Project, Churchill et al. 4698 (CAS), Knapp & Vodicka 5671 (CAS, MO); Chiriqui: highway between
Gualaca and Chiriqui Grande, just south of continental divide trail, Croat 66534 (CAS, US), 10 km N
of Los Planes de Hornito, near IRHE Fortuna Hydroelectric Project, Antonio 5180 (CAS), Churchill
5251 (CAS), Correa 2143 (US), Gordon 300 (CAS), Knapp 5023 (CAS, MO), Mendoza 327 (US),
Valdespino et al. 558 (PMA). San Blas: Cerro Habu, 1,400—2,500 ft., Sytsma et al. 2665 (MO),
Sytsma et al. 2745 (CAS, MO). Darién: Cerro Pirre, 750-1560 m elev.; Gentry & Clewell 6956 (MO),
Duke 6083 (GH, MO), Duke 5342 (MO), Duke 6597 (MO), Duke & Ellias 13815 (MO), Duke &
Ellias 13794 (MO), Bristan 539 (MO), Croat 37870 (MO), Folsom 4248 (CAS, MO), Folsom et al.
4468 (CAS, MO, PMA); ridge from Altos de Nique to Cerro Pirre, Croat 37842 (MO), Cana and vi-
cinity, 2000-6500 ft., Williams 812 (US, NY); Cuasi-Cana trail between Cerro Campamiento and La
Escalera to “Paramo,” east of Tres Bocas, Kirkbride & Duke 1344 (MO, NY); Crest Cana Cuasi Trail,
Real District, Terry & Terry 1543 (F, GH).

DISCUSSION. — Monolena primuliflora Hook.f. was the first species of Monolena described, that
having been done in 1870. Hooker’s (1870) descriptions and plate of M. primuliflora in the Botanical
Magazine and the plate and description of Bertolonia primuliflora in the Floral Magazine (Dombrain
1870) were apparently published at about the same time. The Hooker description was published in
February while the Dombrain article cannot be specifically dated. Both descriptions are discussed in
the March 1870 volumes of Gardener’s Chronicle (Anonymous 1870, Anonymous 1870a).

The description and plate in the Botanical Magazine were prepared from plants cultivated in Eng-
land from plants originally introduced by Wallis from the Rio Zamora, Ecuador (Anonymous 1870,
Anonymous 1870a). No herbarium voucher for these is known. The specific epithet has usually been
spelled “primulaeflora.” This is corrected here to “primuliflora,” in accordance with Art. 60.8 of the
ICBN (Greuter et al. 2000)

This species is distinguished by a combination of characters, the most important being leaves that
are ovate to elliptic, cuneate to obtuse or rarely subcordate at the base, and nerved to only slightly
plinerved (except for plants from the Talamanca Mountains of Costa Rica and Panama with strongly
plinerved leaf blades) and an inflorescence typically bearing 3—6 flowers. Variation within this spe-

114 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES
Volume 53, No. 9

cies is considerable, particularly in the Talamanca Mountains, at lower elevations along the Pacific
coast of Colombia and Ecuador, and in Acre, Brazil.

In M. primuliflora foliar sclereids are typically moderately abundant in the lower hypodermis.
However, plants at low elevations along the Pacific lowlands of Colombia have a very low density of
foliar sclereids. Apparently following this pattern, plants from low elevations (300-400 m) in Costa
Rica have membranous leaves, suggesting a low density of foliar sclereids.

8. Monolena trichopoda R. H. Warner, sp. nov.

TYPE: PANAMA, Veraguas, NW of Santa Fe, 2.7 km from Escuela Agricola Alto de Piedra on
road to Calovébora; on rocks along stream; corolla white, flushed with pink; 30 March 1975; S. Mori
& J. Kallunki 5335 (holotype: MO!; isotype: US!).

Laminae elliptico-lanceolatae, basi cuneata usque obtusa, valde plinerves; petioli setosi, setae
ephemerae in series duas; inflorescentia floribus 3—S (—8); bractae florales ellipticae usque anguste
obovatae; petala 1 1—16 (—18) mm longa; appendices ventrali linguiformi staminum antisepalum.

Epiphytes on low branches, and on logs and rocks. Dried rhizomes 1—4 x 0.3—1.0 cm. Leaves
membranaceous when dry, puberulent throughout, the hairs ca. 0.1 mm long; juvenile leaves occa-
sionally bearing hairs 2.0—2.5 mm long on the adaxial surface; petioles purple, 2-12 (—15) cm long,
bearing two rows of setae 0.9—2.5 (—3.0) mm long along the lateral ridges; upper blade surface green;
lower surface purple, often turning red in the areoles and yellow-green along the primary and the sec-
ondary veins on drying; blades 7—22.5 x 2.0—10 cm, elliptic-lanceolate, cuneate to obtuse at the base,
acute to subacuminate at the apex, strongly 3—5 (—7, rarely 9) plinerved, the innermost pair of primary
veins diverging from the midrib in an alternate to rarely opposite arrangement 1/5—1/3 of the distance
to the apex; primary veins prominent below; margin entire. Inflorescence bearing 3—5 (—8) flowers;
peduncle and axis of the inflorescence purple, together 5.5—15 (—19) cm long in flower and 18—24
(41) cm long in fruit when measured to the furthest pedicel, green or purple, scantily puberulent with
hairs 0.1—0.2 mm long; bracts and scales of the inflorescence caducous, translucent white, petaloid,
7-12 x 3-9mm, elliptic to narrowly obovate, obtuse to rarely emarginate at the apex,
concavo-convex, entire; scales 2 or sometimes apparently only one; lower and lateral floral bracts
each one per flower. Pedicels 2.0-3.5 mm long. Hypanthium green, 3.0—-3.5 mm long, 3.5—4.0
(—5.0) mm wide at the torus. Calyx tube 0.2—0.3 (—0.5) mm long; calyx lobes green or pink to purple,
1.5-2.0mm long, 2.0-2.5mm wide at the base, triangular, rounded at the apex. Pedicels,
hypanthium, and the margin and both sides of the calyx scantily puberulent, the hairs 0.1—0.2 mm
long. Petals white at the base and pink toward the apex, | 1-16 (—18) x 6-10 mm, obovate, rounded at
the apex, entire. Antesepalous stamens: filaments 4.1—5.0 (—5.5)mm long; thecae yellow,
2.0—2.5 mm long; connective extended |.6—1.8 mm between the thecae and the insertion of the fila-
ment, bearing a dorsal tubercle ca. 0.1 mm long, and a tongue-shaped ventral appendage 0.9—1.3 mm
long. Antepetalous stamens: filaments 3—4 mm long; thecae yellow, 2.0—2.2 (—2.5) mm long; connec-
tive yellow, extended 0.8—1.0 mm between the thecae and the insertion of the filament, the ventral ap-
pendage 0.2—0.4 mm long. The connective in both whorls of stamens bending in late anthesis to bring
the thecal pore into contact with the stigma. Ovary ca. 0.3 mm long; style 6.0—6.5 (7.0) mm long, and
1.4—1.6 mm wide where expanded. Fruit 1.0—1.3 mm wide. Seeds ca. 0.7 x 0.5 x 0.4 mm.

DISTRIBUTION AND PHENOLOGY. — This species of Monolena is endemic to Panama where it is
found in the provinces of Veraguas and Coclé from the region of the continental divide north to the Ca-
ribbean lowlands. It is locally common along rocky streams between 400—1200 meters. Fertile speci-
mens have been collected every month except April, including flowers and fruits in most months.

WARNER: MONOLENA IN CENTRAL AMERICA 115

PARATYPES. — PANAMA. Coclé: El Copé, vicinity of sawmill, north across continental divide,
and Alto Calvario, Antonio 2076 (CAS, PMA), Croat 44610 (MO), Croat 68800 (MO), Galdames et
al. 1070 (PMA), Hamilton & Davidse 2770 (MO), Hammel 801 (CAS), Folsom 1595 (MO), Folsom
3268 (CAS), Folsom 4118 (CAS, MO), Folsom & Page 5966 (CAS), Folsom & Collins 6527 (MO),
Folsom et al. 7089 (CAS), Read et al. 81-18 (US), Sytsma 1922 (CAS), Sytsma & Anderson 4538
(MO), Warner 420 (MIN); Rio Blanco, 2 hour walk down Atlantic slope from sawmill at El Copé, R.
Dressler 5646 (PMA), Warner 419 (MIN); Rio Tife, base of Cerro Tife, S. Knapp 3708 (MO), S.
Knapp 3719 (CAS, MO); Veraguas: road from Agricultural School Alto de Piedra to Calovébora:
Alto de Piedra, Tam 27 (MO, PMA); Rio Primero Brazo, ca. 2.6 km beyond the school, 7. Antonio
2037 (CAS, MO), Croat 23179 (MO), Croat 25439 (MO), Croat 25449 (MO), Croat & Folsom
34121 (MO), Mori & Kallunki 4759 (MO), Warner 418 (MIN); Rio Segundo Brazo, Mass & Dressler
1633 (MO, US); 8.8-11 km from the school, Mori & Kallunki 3206 (PMA) Mori & Kallunki 6203
(MO); Rio Dos Bocas, Croat 27406 (MO, PMA) Croat 27506 (MO), Mori et al. 3861 (MO); Guabal,
between the Continental Divide and Rio Calovébora, Dressler 4719 (MO, PMA, US).

DISCUSSION. — Monolena trichopoda is recognized by the two rows of cilia on the petioles, the
leaf blades strongly plinerved with the margins entire, the small number of scales on the inflorescence,
and the tongue-shaped ventral appendage on the connective of the antesepalous stamens. On the one
hand, these plants share certain features with those of the Cordifolia complex, such as small stature,
absence of foliar sclereids, and bending of the connective in late anthesis to bring the thecal pore into
contact with the stigma. On the other hand, the blades are lanceolate to elliptic and strongly plinerved,
characters not found in the Cordifolia complex. Several specimens from Bocas del Toro, Panama
(Santamaria & Lara 952, 964 and 1078, all at PMA) are related to Monolena trichopoda, but are left
unassigned pending further study. The petioles on these specimens have a dense pubescence that is
perhaps unique for the genus.

ETYMOLOGY. — The specific epithet refers to the ciliate petiole, a character that is unique to this
species of Monolena.

ACKNOWLEDGMENTS

I wish to thank Dr. Thomas Morley who was my teacher and mentor. Dr. Frank Almeda and Dr.
John Wurdack freely shared their knowledge about the Melastomataceae. Cathy Pasquale drew the il-
lustrations; Juan M. Ochoa made the distribution map. Many present and former staff members of the
Missouri Botanical Garden assisted with field studies; Thomas Antonio and Barry Hammel in partic-
ular provided pickled flowers and live plants of Monolena. | am grateful to the curators of the follow-
ing herbaria who loaned specimens for study: BM, BR, CAS, F, GH, K, L, MO, NY, P, PMA, S, US,
and WIS.

Field studies were supported by: The Alexander Bache Fund, National Academy of Sciences;
Sigma XI, Grants-in-Aid of Research; and from the University of Minnesota: The Wilkie Fund of the
James Ford Bell Museum of Natural History, Anderson Fellowship of The Graduate School, and the
Butters Fund of the Department of Botany. Special assistance provided by Dr. Donald and Elizabeth
Lawrence is gratefully acknowledged.

LITERATURE CITED

ALMEDA, F. 1978. Systematics of the genus Monochaetum (Melastomataceae) in Mexico and Central America.
Univ. Calif. Publ Bot. 75:1—134.

ANONYMOUS. 1870. Gardener’s Chronicle. No. 10:309.

ANONYMOUS. 1870a. Gardener’s Chronicle. No. 11:344.

COGNIAUX, C. A. 1891. Mélastomacées. In Monographiae Phanerogamarum, Vol. 7. A. and C. de Candolle, eds.
G. Mason, Paris. 1256 pp.

116 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES
Volume 53, No. 9

GLEASON, H. A. 1958. Melastomataceae. /n Flora of Panama, R. E. Woodson Jr. and R. W. Shery, eds. Ann.
Missouri Bot. Gard. 45:203-305.

. 1930. Studies on the flora of northern South America - XIV. Melastomaceae from Colombia and Ec-
uador. Bull. Torrey Bot. Club 57:63-—75.

GREUTER, W. et al. 2000. International Code of Botanical Nomenclature. Regnum Vegetabile (Vol. 138).
Koeltz Scientific Books, K6nigstein, Germany.

HEEL, W. A. VAN. 1958. The pistil of Bertolonia marmorata Naud. (Melastomataceae). Blumea, Suppl.
IV:144-148.

Hooker, J. D. 1867. Melastomataceae. Pp. 721—952 in Genera Plantarum, Vol. 1, G. Benthum and J. D.
Hooker eds. L. Reeve and Co., London.

MACBRIDE, W. 1941. Flora of Peru. Field Mus. Nat. Hist., Bot. Ser. 13(1V—I):1—565.

SOLT, M.L. AND J.J. WURDACK. 1980. Chromosome numbers in the Melastomataceae. Phytologia
47:199-220.

STANDLEY, P. C. AND L. O. WILLIAMS. 1963. Melastomataceae. /n Flora of Guatemala. Fieldiana, Bot.
24:470-S25.

Tost, J. 1970. Mapa Ecologico de Panama. Programma de las Naciones Unidas para el Desarrollo. Rome.

TRIANA, J. J. 1871. Les Mélastomacées. Trans. Linn. Soc. 28:1—188.

URIBE, L. 1960. Sertula Florae Colombiae, V. Caldasia 8:319—324.

. 1971. Sertula Florae Colombiae, XII. Caldasia 11:81—91.

. 1971. Nueva especie colombiana de Melastomataceas. Mutisia 46:1—3.

WHIFFIN, T. AND S. TOMB. 1972. The systematic significance of seed morphology in the neotropical capsular
fruited Melastomataceae. Amer. J. Bot. 59:41 1-422.

WURDACK, J. J. 1980. Melastomataceae. /n Flora of Ecuador 13:1—406, G. Harling and B Sparre, eds. Univer-
sity of Géteborg; Riksmuseum, G6teborg, Sweden.

_ a
7) ,
‘—

7
, y : > ? 7 : -
oes Ove er iibea Ta
: ; ‘ : . » te - a
Cup sirta Al Cl, wabeayie, de lett oma Du. a
tee ee oi -Oti > 7 es
—s Si indo da ter areata ruth Areas ty °f Letras i

gate, , Pra ee ON 9) 7 - 7
1 ‘Ste bal wwe Th @ liege ia us) Wut ind rated Lena
2 Pateiands tod Fe nine . maa _
i * Ce de | Prey [iP A ; m, wats)

4) «68

on = Se. 7 oat
7 - 7 j - _
[ a ‘ 7 ot -
-
7 7
el :
¥
?
——
¥*

PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES

Volume 53, No. 10, pp. 117-149, 63 figs. August 22, 2002

A Revision of the African Spider Genus Raecius Simon, 1892
(Araneae, Zorocratidae)

Charles E. Griswold
Schlinger Curator of Arachnida, Department of Entomology, California Academy of Sciences,
Golden Gate Park, San Francisco, California 94118
and
Research Professor of Biology, San Francisco State University
email: cgriswold@calacademy.org

The tropical African spider genus Raecius Simon, 1892 is defined and revised and a key to
the five valid species is provided. Raecius congoensis, R. jocquei and R. scharffi new species
are described. Mnesitheus vittatus Simon, 1907 and M. zoropsides Strand, 1916 are new ju-
nior synonyms of Raecius asper (Thorell, 1899). Raecius aculeatus Dahl, 1901 is excluded
from the genus and is incertae sedis in Zorocratidae.

Raecius are cryptic spiders in their habits and history. They are rare in collections, usually col-
lected singly or a few at a time, and the only series in collections have resulted from mass collecting
through pitfall trapping or concerted efforts to dig them from burrows. Raecius occur in the disjunct,
Afromontane forests (Griswold 1991la; White 1978, 1983) in tropical Africa (e.g., Mt. Cameroon,
Bioko Island, the Eastern Arc mountains of Tanzania) but are not restricted to these habitats, occur-
ring as well in lowland forests and arid areas. In this, the 16th in a series of monographs on spiders oc-
curring in the Afromontane region (Griswold 1985, 1987b, 1987c, 1987d, 1990, 1991b, 1994, 1997a,
1997b, 1998a, 1998b, 2000, 2001; Griswold and Platnick 1987; Griswold and Ledford 2001), I revise
the genus Raecius and discuss its taxonomic placement in the Zorocratidae.

The spiders placed in Raecius have a complex and confused taxonomic history. Indeed, for much
of the 20th century synonyms of the same species were placed in different families, the Tengellidae
and Zoropsidae. Koch (1875:32) described the first Raecius species from a juvenile from Ethiopia (in
what is today Eritrea). He called it Amaurobius crassipes, which he placed in the Agelenidae. Simon
(1892:230) proposed the genus Raecius for Koch’s species and placed it in the Zoropsidae along with
the genera Zoropsis and Zorocrates. Thorell (1899:18) proposed Mnesitheus asper for a female spider
from Mt. Cameroon that he placed in the Dictynidae. Simon (1903:975) listed Mnesitheus in the
Dictynidae, and in 1907 described another species M. vittatus from Bioko Island (Simon 1907:227).
In 1916 Strand described a male spider from Mt. Cameroon as Mnesitheus zoropsides (Strand
1916:140). Dahl (1901a, 1901b, 1913) treated Raecius and Mnesitheus in the Zorocratidae, but this
placement was not followed by subsequent workers. Petrunkevitch (1928) listed Raecius in the
Zoropsidae and Mnesitheus in the Tengellidae, a placement followed by Roewer (1954) and Bonnet
(1957, 1958). It was not until Lehtinen’s comprehensive study of type specimens of cribellate spiders
that the synonymy of these genera was recognized (Lehtinen 1967:250, 262). Lehtinen synonymized
Mnesitheus with Raecius, and transferred the latter genus into his Miturgidae, Uliodoninae. I under-
took the first quantitative phylogenetic analysis of exemplars for taxa placed in the Tengellidae,

118 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES
Volume 53, No. 10

Miturgidae, Zoropsidae and Lycosoidea (Griswold 1993). This analysis treated taxa that had one or
more classical (i.e., grate-shaped tapetum) or newly discovered characters (e.g., male tibial crack,
oval calamistrum, and/or interlocking lobes on the tegulum and subtegulum of the male pedipalpal
bulb). Significant results were that Raecius formed a clade along with the other former miturgid gen-
era Campostichomma, Zorocrates, Zorodictyna, and Uduba. These taxa were not closely related to
Miturgidae (represented in that study by Uliodon tarantulinus) but no family transfer was made.
Within this clade the entire cribellum suggested that the Malagasy genus Zorodictyna and African
Raecius are sister groups. Griswold, Coddington, Platnick, and Forster (1999) examined an even
broader array of araneomorph taxa. Their phylogenetic analysis suggested that the former miturgid
genera Raecius and Uduba (and by implication, Campostichomma, Zorocrates, and Zorodictyna)
comprise the family Zorocratidae Dahl 1913. Synapomorphies of Zorocratidae are the male tibial
crack, clumped cribellar spigots (Figs. 19,21), and a male pedipalpal tibia ventroapical process (Figs.
22,38):

Comparison to Zorodictyna and more distantly related zorocratids suggests several synapo-
morphies for Raecius. The long, narrow carapace (Fig. 1) contrasts with the oval carapaces of other
zorocratids. The flattened, ribbon-like embolus (Figs. 37, 51) and blunt embolic tip (Figs. 40, 62) may
be additional Raecius synapomorphies as other zorocratids have cylindrical or attenuate emboli
(Griswold 1993, figs. 11, 13, 20, 34). Phylogenetic resolution within Raecius is difficult because R.
congoensis is known only from females and R. scharffi only from males, but at least the enlarged RTA
may be a synapomorphy uniting R. asper and R. jocquei.

The sister group relationship of Raecius and the Malagasy Zorodictyna 1s of particular interest,
being one of at least five documented cases of Madagascar-tropical African montane, vicariant sister
group relationships (Griswold 2000). Four other parallel patterns occur within Cyatholipidae (within
Ulwembua and between Alaranea and Scharffia) and Phyxelididae (within Phyxelida and between
Ambohima and Kulalania). This corroborates the hypothesis in Griswold (1991a) suggesting that
Madagascar is related to areas in tropical East and South Africa. This tropical A frican-Madagascar
vicariance pattern parallels that suggested for lemurs (Yoder et al. 1996) and tenrecs (Asher 1997).
Recent molecular clock evidence suggests that the mammalian disjunctions may be Eocene in age
(Yoder et al. 1996), and a land bridge along the Davie Fracture Zone has been postulated as a path for
dispersal from A frica to Madagascar at this time (McCall 1997). The replicated biogeographic pattern
among spiders may also result from dispersal during the early Tertiary.

METHODS

Species descriptions represent a single individual of each sex that is noted at the head of each de-
scription. The section on variation represents the variability in several individuals of each sex chosen
to represent the full range of overall size. All measurements are in millimeters. The length and width
of the cephalothorax and abdomen were measured from above and represent maximum values,
chelicerae were excluded from cephalothorax length but included in measurements of total length;
carapace height was measured from the side and represents the maximum height; the ocular area was
measured from above with the dimensions of the ocular quadrangle being measured from the outer
margin of the eyes; the posterior eye row was measured from above and the anterior eye row was mea-
sured from in front; eye interdistances are abbreviated, e.g., AME-AME = distance between anterior
median eyes, ALE-PLE = distance between lateral eyes; clypeal height represents the distance from
the median margin of the clypeus to the lower margin of the AME; cheliceral length comprises the dis-
tance from the median margin of the clypeus to the apex of the paturon (not including the fang);
lengths of leg segments were measured along the dorsal margin as in Griswold (1987a). Spines are re-
ported for the dorsal (d), prolateral (p), ventral (v), and retrolateral (r) surfaces of the legs and
pedipalpi and are noted from the proximal to distal ends of each segment. Abbreviations for the ocular

GRISWOLD: AFRICAN RAECIUS SPECIES 119

region (AME, ALE, PME, PLE, AER, PER) and legs are standard for the Araneae. A list of abbrevia-
tions used to refer to special morphological features is presented in Table |. Label data are quoted ver-
batim in the listings of material examined.

Tracheae were examined by first dissecting off the dorsal cuticle of the abdomen, digesting away
nonchitinous tissue in 10 percent potassium hydroxide (KOH) either warmed beneath a desk lamp for
72-120 hours or boiled for 5—10 minutes, and then staining the tracheae and other chitinous structures
in Chlorazol Black. Male pedipalpi were expanded by immersing them overnight in a weak, watery
solution of KOH and transferring them to distilled water where expansion continued. Pedipalpi were
transferred back and forth between KOH and distilled water until expansion stopped. Female genitalia
were excised from the abdomen, cleaned in a solution of trypsin and water, and illustrated. Pedipalpi
and vulvae were examined in alcohol and in lactic acid. Subsequently, to allow examination of inter-
nal structures, vulvae were bleached through brief immersion at room temperature in a 5.25% sodium
hypochlorite solution (CLOROX® household bleach). Bleaching provided rapid, excellent clearing
of even the most heavily sclerotized genitalia with no apparent distortion of features. Vulvae were
then stained lightly with Chlorazol Black, temporarily mounted in lactic acid, and examined with a
compound microscope. Terminology for parts of the vulva follows Sierwald (1989). Prior to exami-
nation with a Hitachi S-520 Scanning Electron Microscope all structures were cleaned in an ultrasonic
cleaner: pedipalpi and female genitalia were critical point dried, other structures were air dried.
Spinnerets were prepared by first causing full extension by squeezing the abdomen with forceps, lock-
ing the forceps in position with a paper clip, and transferring the spider and forceps to absolute ethanol
for 24 hours. The whole abdomen was then cleaned with an ultrasonic cleaner, critical point dried, and
mounted on a stub using Elmer’s Glue. Tapeta in the indirect eyes were examined in living or freshly
dead specimens as in Griswold (1993). Specimens were examined with a Wild MSA, Olympus SZH
or Leica MZAPO stereomicroscope, and/or a Nikon compound microscope modified for 3 dimen-
sional imaging by SL3D, Inc., and drawings were made with a mounted phototube (camera lucida).

TABLE |. Abbreviations for morphological terms used in text and figures.

AC — aciniform gland spigot(s)

AER ~— anterior eye row

ALE — anterior lateral eyes

ALS ~— anterior lateral spinneret

AME ~ anterior median eyes

AN — anelli

C — conductor

BS — base of spermatheca

CD — copulatory duct

CY — cylindrical gland spigot(s)

DTA — dorsal apophysis of tibia

E — embolus

EF — epigynal fold

FD — fertilization duct

HS — head of spermatheca

ITC — inferior tarsal claw

LL — lateral lobes of epigynum

MA - median apophysis of tegulum
MAP — major ampullate gland spigot(s)
mAP — minor ampullate gland spigot(s)

ML — median lobe of epigynum

MS — median sector of epigynum

N — nubbin

OAL — length of ocular area

OQA — ocular quadrangle, anterior
OQL — ocular quadrangle, length
OQP — ocular quadrangle, posterior
P — petiole

PER — posterior eye row

PI — piriform gland spigot(s)

PLE — posterior lateral eyes

PLS — posterior lateral spinneret
PME — posterior median eyes

PMS — posterior median spinneret
PsFl — pseudoflagelliform gland spigot(s)
RTA ~- retrolateral apophysis of tibia
ST — subtegulum

STP — sclerotized tegular process

T — tegulum

VTA — ventral apophysis of tibia

120 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES
Volume 53, No. 10

Throughout the text references to figures from other papers are noted in lower case (fig.), whereas ref-
erences to figures accompanying this paper are capitalized (Fig.).

NATURAL HISTORY

Raecius have been observed in the field in Cameroon, on Bioko Island, Equatorial Guinea and in
Tanzania. They occur in lowland and montane forest (Fig. 26) and in montane grassland (Fig. 27) and
at least R. asper occurs in both montane forest and montane grassland. The type of R. crassipes was
taken in a dry riverbed in northern Eritrea, a habitat much drier than that of other Raecius species.
They are burrowing or tube building spiders that line their burrows with silk and provide the entrance
with a collar of cribellate silk (Fig. 25). Burrows of R. asper from Mt. Cameroon were 6—8 cm deep,
and straight or slightly curved. In forest, burrows are typically made in rotting fallen logs, or silken
tubes or tunnels may be made in moss on tree trunks. In grassland the burrows are in turf and may be
abundant. In burned grassland in January on Mt. Cameroon (Fig. 27), when the burrows were most
conspicuous, 3 person-hours of collecting revealed more than 70 individuals. Many adult females at
this locality had egg sacs or young in the burrow. These egg sacs were thickly wrapped with silk and
covered with dirt or other debris. Populations from forest and grassland on Mt. Cameroon differed in
phenology. Two egg sacs collected in mist forest at 1425 m had only eggs within (37 and 41 eggs),
whereas sixteen egg sacs collected during the same week from grassland at 2050 m had juveniles or
were already empty. Of these sixteen, nine were empty and seven had 23 to 55 (average 42) juveniles.
A female from Mt. Kupé had an egg sac containing approximately 110 eggs. One to two egg sacs
could be found in a female’s tube, with one sac always more advanced phenologically than the other.
A captive individual of R. scharffi captured prey with a grab and bite. Prey was carried in the
chelicerae and was not wrapped. Captive individuals of both R. asper and R. scharffi carded cribellate
silk using legs IV braced against each other and moved simultaneously. This “mobile leg [V” carding
is typical of entelegynes (Eberhard 1988, Griswold et al. 1999). Cribellate silk has both axial lines and
reserve warp.

TAXONOMY

Raecius Simon, 1892

Raecius Simon, 1892:230 (type species Amaurobius crassipes L. Koch 1875:32, from Eritrea, in BMNH, exam-
ined, described in Agelenidae). Petrunkevitch 1928:146. Roewer 1954:1283. Bonnet 1958:3846. Lehtinen
1967:250 (transferred to Miturgidae). Brignoli 1983:544. Platnick 1989:432; 1993:598; 1997:682; 2002.
Griswold et al. 1999:59 (transferred to Zorocratidae).

Mnesitheus Thorell, 1899:18 (type species Mnesitheus asper Thorell, from Cameroon, in RMS, examined, de-
scribed in Dictynidae). Simon 1903:975. Petrunkevitch 1928:91. Roewer 1954:1379. Bonnet 1957:2968.
Synonymized by Lehtinen 1967:250.

DIAGNOSIS. — Zorocratid spiders with entire cribellum (Figs. 18-19) and oval calamistrum
(Fig. 8) that differ from Zorodictyna in having feathery hairs (Figs. 6, 9, 43) and lacking a dorsobasal
projection of the cymbium, and that differ from all other zorocratids in the flattened, ribbon-like
embolus (Figs. 37, 51), blunt embolic tip (Figs. 40, 62), and elongate carapace with length 1.47 to 1.90
times width (Fig. 1).

DESCRIPTION. — Medium sized to large spiders, total length 4.68—15.73; sexual dimorphism
slight, male with relatively longer legs. Carapace elongate-oval in dorsal view (Figs. 1, 24), length
1.47 to 1.90 times width, domed (Fig. 2), height 0.38 to 0.74 times width; fovea linear, deep. Ocular
area broad, PER width 2.344.87 times OA length; ocular quadrangle rectangular to trapezoidal, in
most specimens widest behind, OQP/OQA 0.93—1.36; eight eyes in 2 nearly straight rows, indirect

GRISWOLD: AFRICAN RAECIUS SPECIES 121

eyes with canoe-shaped tapetum. Clypeus low, height 0.84—2.06 times AME diameter. Chelicerae
stout with large boss (Fig. 2), length 6.31—12.00 times clypeal height, pro- and retromargins of fang
furrow in most specimens with 3 teeth. Sternum length 1.3 1—1.88 times width; labium long, with basal
notch, length 1.19-1.67 times width; pedipalpal coxal endites nearly parallel (Fig. 3), length
1.94-3.00 times width, serrula teeth in single row along outer margin. Leg formula 1423 or 4123, ratio
of femur I length/carapace width | .32—2.84 in male, 0.97—1.87 in female; integument finely wrinkled,
with plumose and feathery setae (sensu Lehtinen 1975) on legs and body (Figs. 6, 9, 43-44); male leg
tibiae with basal crack (Griswold 1993, figs. 3, 4); trochanters with broad, shallow notch. Spination,
male basic pattern, comprises: pedipalpus: femur d0-1-1-1, r0-0-0-1; leg I: femur d1-0-0, tibia
v2-2-2-2, p0-1-1-0, r0-1-1-0, metatarsus v2-2-1, pl-1-1, rl-1-1; leg I: femur d1-0-0, tibia v2-2-2,
p0-1-1-0, r0-1-0, metatarsus v2-2-1, p1l-1-2, r0-1-0; leg III: femur: d1-0-1, tibia: vl-1-2, pl-1-0,
d0-1-0, rl-1-0, metatarsus vl-1-2, pl-1-2, d0-1-0, rl-1-2; leg IV: femur: d1-0-0, tibia v1-2-2,
r0-1-1-0, metatarsus: v1-1-2, p0-0-2, r0-0-2. Raecius jocquei has more dorsal spines on femur and
tibia land R. scharffi has fewer lateral spines on tibia and metatarsus I and fewer ventral spines on tibia
II. Female basic spination pattern: pedipalpus: femur d1-1-1, tibia p0-1-0, tarsus p1-1, vO-1; leg I: fe-
mur: d1-0-1, tibia v2-2-2-2, metatarsus p0-0-2, v2-2-2, r0-0-2; leg II: femur: d1-0-1, patella p1, tibia
pl-0-1, vl-1-2, metatarsus p1-0-2, v2-2-1, r0-0-2; leg III: femur d1-0-0, patella pl, tibia d0-1-0,
pl-0-1, vl-1-2, rl-0-1, metatarsus p 0-1-1-2, v2-2-2, rl-1-2; leg IV: femur d1-0-0, tibia p1-0-0,
vl-1-2, metatarsus p0-0-2, v1-1-2, r0-0-2. Femoral dorsal, patellar prolateral, and lateral spines of the
tibiae and metatarsi may be fewer or more than this pattern. Calamistrum rectangular-oval, arising ba-
sally and extending for 2/3 length segment, attenuate distally (Figs. 6, 8). Trichobothrial pattern: ab-
sent from leg femora and patellae, tibiae with a dorsobasal group, few prolaterals, and a retrolateral
row extending to apex, metatarsi with a dorsal, irregular row, this row restricted to apical half of ['Vth,
tarsi with dorsobasal row that divides into 2-3 rows apically, pedipalpal tibiae with pro- and
retrolateral rows, in males extending dorsad and retrolaterad of RTA, apparently absent from tarsi of
most specimens, although a female of R. jocquei has a single dorsal trichobothrium on one pedipalpal
tarsus; trichobothrial base with transversely-ridged hood (Figs. 4, 7). Tarsal organ median to basal,
capsulate, orifice simple, oval to round (Figs. 5, 7). Male scopulae beneath tarsi I-IV, female scopulae
beneath all tarsi and at least metatarsi I and II (Fig. 1), scopulae extend distad of tarsal apices, may ob-
scure claws; dorsal scopulate patch on cymbium absent (Fig. 53). Preening combs absent. Superior
tarsal claws pectinate, those of legs I and II with 8-12 teeth, those of legs III-IV with 3-7 teeth, infe-
rior tarsal claws (ITC) simple, large or reduced to nubbins or absent; female pedipalpus claw
pectinate, with 3—10 teeth; claw tufts absent. Male pedipalpal tibia with retroapical (RTA) and ventral
(VTA) apophyses (Figs. 22, 38), dorsal apophysis (DTA) present (R. scharffi: Fig. 55) or absent.
Pedipalpal tarsus (Figs. 22, 23) with large, triangular petiole (P) attached to alveolus; subtegulum
(ST) cup-shaped, with 4—5 anelli (AN), prolateral margin with lobe that articulates with correspond-
ing lobe on tegulum in unexpanded bulb (Fig. 23); tegulum (T) U-shaped, simple; firmly attached
embolus (E) arising on prolateral side of tegulum, embolus flattened and ribbon-like, may have
groove or fold along outer margin (Fig. 51); median apophysis (MA) flexibly attached; conductor (C)
hyaline, originating from retroapex of T, cup or fan-shaped (Figs. 29, 45, 55); tegulum with no (R.
scharffi), one (R. asper) or two (R. jocquei) sclerotized processes (STP); fundus in subtegulum, reser-
voir and ejaculatory duct simple, without loops or switchbacks, spiraling around outer margin of T.
Abdomen oval, without scuta (Figs. 1-2). Respiratory system comprising two anterior book lungs and
a single, small spiracle just anteriad of cribellum leading into four simple tracheal tubes confined to
abdomen (examined in juvenile R. asper from Mann’s Spring). Epigynum (Fig. 41) divided by
epigynal folds (EF) into median sector (MS) and lateral lobes (LL); median sector with transverse
lobe (ML), variously shaped; LL tooth present or absent; vulva (Figs. 42, 58; Griswold 1993, figs. 29,
30) with copulatory ducts (CD) anteromedian, short, slender, entering spermathecal head (HS) later-
ally or ventrolaterally; HS small, hemispherical; base of spermatheca (BS) simple, short, fertilization

122 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES
Volume 53, No. 10

duct (FD) posterior. Cribellum wide, short, width greater than 3 times length, spinning field entire
(Figs. 18-19), cribellar spigots arranged in longitudinal linear groups (Figs. 19, 21). Six spinnerets
(Fig. 18), anteriors (ALS) and posterior laterals (PLS) two-segmented, posterior medians (PMS)
one-segmented; ALS conical, PMS and PLS cylindrical, distal segment of PLS domed, short, less
than 1/3 length basal segment. Female (examined in R. asper and R. jocquei) with ALS (Fig. 11) hav-
ing two large major ampullate gland spigots (MAP) on mesal margin and oval field of more than 30
piriform gland spigots (PI); PMS (Fig. 12) with one large minor ampullate gland spigot (mAP) near
anterior margin and large central nubbin, several aciniform gland spigots (AC), with 4-5 large cylin-
drical gland spigots (CY) with long conical bases and long slender shafts on posterior surface of
spinneret extending to base; PLS (Figs. 13, 20) with 4-6 large CY on margin of AC spinning field, ap-
parent apical pseudoflagelliform gland spigot (PsFl) present (R. asper: Fig. 20) or not apparent (R.
Jocquei: Fig. 13). Male (examined in R. jocquei: Figs. 14-17) lacking CY, other spigots reduced in
number: only one MAP on ALS accompanied by a nubbin, and apical nubbin on PLS (Fig. 17). Sucha
PLS apical nubbin may replace the PsFI spigot in the male, suggesting that a PsF] may be present (but
undistinguished) in the female of R. jocquei. Male epiandrous spigots absent (Fig. 44).

COMPOSITION. — Five species.

DISTRIBUTION. — Tropical Africa from Cote d’Ivoire on the west across the Congo basin to
Eritrea and Tanzania in the east (Fig. 63).

KEY TO SPECIES OF RAECIUS

1 Dorsal abdominal markings uniform (Koch 1875, pl. 3, fig. 4); leg formula 4123; occurring in semiarid Eritrea
whey eos med Riwretdonie NT Mined Se Saag Se a cl: IE ert CV Sink Mle “peat RR Ries PCRS Get a 5 crassipes (L. Koch)
Dorsal abdominal markings with at least a faint light longitudinal band (Fig. 1); leg formula 1423; occurring in moist

equatonialvAdricay' 3 ef Sc eT Sh. BO SA) OS Od oh) ee rrr p)
2 (ON) pNall eSuer 445 Bee rypsceret retorlieet leopard nc breaep aor Burtt Sect alias Sh dus 39 tcors bat ee 3
SGMAIES Biv cn Syscodergsts oh aseae aT fhe scopes UP ar eiges S leswte ch areteunbanacace, dosoy que &ilpsyceidince go Om ee 5

3(2) Pedipalpal bulb with STP; median apophysis not produced into distal hook; DTA absent; RTA large, length greater

than! O:5itimes length of tibia.) cha f oe Se ean a bee EES A) DP =
Pedipalpal bulb lacking STP, median apophysis produced into distal hook (Figs. 54-56); pedipalpal tibia with DTA;
RTA small, attenuate at apex, less than 0.4 times length of tibia (Fig.55)............. scharffi new species

4(3) Pedipalpal bulb having median apophysis expanded and bilobate at apex and STP curved and narrow (Fig. 47),
pedipalpal tibia with RTA expanded and bilobate at apex, length greater than 0.85 times length of tibia (Fig. 46)
MR ie ieee Ser Re ie SA LY. ta acd ot ste 2! oda id iy eel a ee Jocquei new species
Pedipalpal bulb having median apophysis cup-shaped and STP broad (Fig. 40), pedipalpal tibia with RTA truncate at

Apex, emaqin USO) times lenain ORY) GS co 5 esos ob ee eas ot oe eee cee asper (Thorell)
5(2) Epigynal median lobe concave posteriorly, length less than 1.0 times width ...................... 6
Epigynal median lobe convex posteriorly, narrow, length 1.2—1.7 times width (Figs. 31, 33,35). . . . asper (Thorell)

6(5) Epigynal median lobe deeply concave posteriorly, U-shaped, length 0.80—0.85 times width, lateral lobes with teeth
(Fig. 48)
AL bash ie eerie a seeing te ON Asa Mp ike MaMa Sule t URES BNR eer halt Mi a a Meee Pei. 5: jJocquei new species
Epigynal median lobe weakly concave posteriorly, very broad, length less than 0.55 times width, lateral lobes without
teeth (ISAs) Aver are oe ie Pe ee hd eae Qe hola ee cr 2Ae & adnan) wees ee Se congoensis new species

Raecius asper (Thorell, 1899)
Figures 1—3, 6, 8, 18, 20, 2442, 63

Mnesitheus asper Thorell, 1899:18 (female type specimen from Mapanja, Cameroon, 1884-1885, Valdau and
Knutson, in RMS, examined). Simon 1903:975. Roewer 1954:1379. Bonnet 1957:2968.
Raecius asper, Lehtinen, 1967:262. Platnick 1997:682; 2002.

GRISWOLD: AFRICAN RAECIUS SPECIES 123

Mnesitheus vittatus Simon, 1907:227 (three juvenile syntypes from Moka (sic), Fernando Poo [Bioko Island,
Equatorial Guinea], in MNHN, AR817, examined). Roewer 1954: 1379. Bonnet 1957: 2968. NEW SYN-
ONYMY.

Raecius vittatus, Lehtinen 1967:262. Platnick 2002.

Mnesitheus zoropsides Strand, 1916:140 (male type specimen from Buea, Cameroon, in NHMV, examined).
Roewer 1954:1379. Bonnet 1957:2968. NEW SYNONYMY.

Raecius zoropsides, Lehtinen, 1967:262. Griswold 1993:7. Platnick 1997:682; 2002. Griswold and Ubick
198s 1:

Notes. — Males, females and juveniles have been collected in association on Bioko Island,
Equatorial Guinea. Males correspond in detail to R. zoropsides Strand, juveniles to R. vittatus Simon,
and females to R. asper Thorell. The latter name is the senior synonym. This species was discussed
and illustrated in Griswold (1993:14, figs. 25-27). Data from the holotype of Mnesitheus zoropsides
provided the male Raecius characters in the phylogenetic matrix. This species also provided exemplar
data for Raecius in Griswold et al. (1999).

DIAGNOSIS. — Male with pedipalpal bulb having median apophysis cup-shaped and STP promi-
nent and broad (Figs. 30, 37-40), pedipalpal tibia with RTA truncate at apex, 0.5—0.67 times the
length of the tibia (Fig. 29). Female with epigynal median lobe convex posteriorly, narrow, length
1.2—1.7 times width (Figs. 31, 33, 35, 41).

MALE (Moca, Bioko). — Total length 10.00. Carapace, chelicerae, legs and pedipalpi yel-
low-brown, labium and pedipalpal coxae yellow-brown, lighter at tips, sternum, leg coxae and tro-
chanters yellow-white; carapace with sides of caput and pars thoracica dusky and covered with dark
setae, center of carapace bare and pale; caput dark at anterolateral margins above cheliceral bases;
dark pigment surrounding each eye and extending between ALE and PLE; retrolateral process of
pedipalpal tibia (RTA) dark brown to black; dorsum of abdomen dark gray with narrow central yel-
low-gray longitudinal band, simple anteriorly and scalloped and broken into spots posteriorly; sides
and venter yellow-gray mottled with dark spots; spinnerets yellow-gray. Carapace 5.33 long, 3.24
wide, 1.81 high; clypeus 0.24 high, height 1.04 times AME diameter; ocular area 0.57 long, 1.47 wide,
OAL 1.30 times OQL; ratio of eyes AME:ALE:PME:PLE, 1.14:1.07:1.14:1.00, diameter of PME
0.23: AME interdistance 0.5 times AME diameter, AME-ALE 0.44 times AME; PME interdistance
0.56 times PME, PME-PLE 1.44 times PME; ALE-PLE 0.57 times PLE. Chelicerae 0.98 long. Ster-
num 2.75 long, 1.69 wide; labium 1.20 long, 0.78 wide; pedipalpal coxae 1.73 long, 0.78 wide. Femur
I length 1.49 times carapace width. Spination: basic pattern. ITC I and II absent, III and IV reduced to
nubbins. Leg measurements (Femur + Patella + Tibia + Metatarsus + Tarsus = [Total]): I: 4.83 + 2.05
44,83 -74.15+ 2.18 =[18.04]; I: 3:85+ 1.80 + 3.37 + 3.02 +2.10=[14.14]; IM: 3.07 + 1.46+ 1.804
2.44 + 1.61 =[10.38]; IV: 4.39 + 1.80 + 3.66 + 3.71 + 2.00 =[15.56]; pedipalpus: 2.10 + 0.93 + 0.88 +
[absent] + 1.80 =[5.71]. Pedipalpal tibia with RTA curved, widened at truncate, serrated apex, length
().5—0.67 times that of tibia, VTA cylindrical, tapering to apex, pro- and retroventral margin of tibia at
tibia-tarsus joint produced into small processes. Pedipalpal bulb with median apophysis cup-shaped,
STP prominent and broad, narrowed apically, embolus short and ribbon-like, and conductor small and
narrow (Figs. 28—30, 37-40; Griswold 1993, figs. 25—27).

Variation (N=5). Total length 8.71—12.14; ratios of carapace length/width 1.59—1.65,
height/width 0.44—0.56, ratios of PER/OQP 2.37—2.90, PER/OAL 2.59-3.10, OQP/OQA 1.06-1.21,
ratios of clypeal height/diameter AME 0.88—1.50, cheliceral length/clypeal height 7.68—10.73, ratio
of length femur I/carapace width 1.34—1.40. Markings of abdomen range from a faint anterior longitu-
dinal light mark and uniformly pale sides to a bold longitudinal light mark breaking into chevrons pos-
teriorly with sides mottled. ITC I and II may be reduced to nubbins or absent.

FEMALE (Mann’s Spring, Cameroon). — Total length 12.27. Carapace red-brown, darkest ante-
riorly and on sides of pars cephalica, palest around thoracic fovea, sides covered sparsely with dark
scales; chelicerae dark brown except boss red; labium and pedipalpal coxae red-brown, lighter at

124 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES
Volume 53, No. 10

bases and tips; sternum and leg coxae yellow-brown, legs I and II shading to dusky red-brown from
basal third of tibiae to apex of tarsi; abdomen marked as in male (Figs. 1—3). Carapace 6.29 long, 3.90
wide, 2.19 high; clypeus 0.35 high, height 1.75 times AME diameter; ocular area 0.62 long, 1.88 wide,
OAL 1.32 times OQL; ratio of eyes AME:ALE:PME:PLE, 1.00:1.28:1.21:1.07, diameter of PME
0.24: AME-AME interdistance 0.86 times AME diameter, AME-ALE 1.07 times AME; PME-PME
interdistance 0.70 times PME, PME-PLE 1.82 times PME; ALE-PLE 0.93 times PLE. Chelicerae
2.94 long. Sternum 2.82 long, 1.92 wide; labium 1.41 long, 0.98 wide; pedipalpal coxae 2.16 long,
0.90 wide. Femur I length 1.05 times carapace width. Spination: basic pattern. ITC I and II reduced to
nubbins, [I small, IV large. Leg measurements (Femur + Patella + Tibia + Metatarsus + Tarsus = [To-
tal]): 1: 4.10.4 2.05 +3.51 + 2.88 +2.20=([14.74]; I: 3.80 + 1.85 +2.73 + 2.29 1.95 =(l2362) 5 ie
2.93 + 1.61 + 1.46 + 1.85 + 1.32 = [9.17]; IV: 4.00 + 2.00 + 2.93 + 2.88 + 1.56 =[13.37]; pedipalpus:
2.10+ 1.07+ 1.12 + [absent] + 1.61 =[5.90]. Epigynum with median lobe convex posteriorly, narrow,
length 1.3 times width, lateral lobes contiguous posteriorly, without teeth (Fig. 31). Vulva with
spermathecal head oval (Fig. 32).

Variation (N= 11). Total length 10.85—14.42; ratios of carapace length/width 1.47-—1.53,
height/width 0.53—0.73, ratios of PER/OQP 2.56—3.64, PER/OAL 2.89-4.87, OQP/OQA 1.08-1.36,
ratios of clypeal height/diameter AME 1.48—2.06, cheliceral length/clypeal height 7.32—9.41, ratio of
length femur I/carapace width 0.97—1.14. Markings of abdomen range from a faint anterior longitudi-
nal light mark and uniform sides to a bold longitudinal light mark breaking into chevrons posteriorly
with sides mottled (Figs. 1, 2). Patellar spines present or absent. Scopulae restricted to tarsus and
metatarsus I and tarsus II or extending onto tibia I and metatarsus I]. Epigyna with lateral lobe teeth
present (Figs. 33, 35) or absent (Fig. 31), median lobe length 1.2—1.7 times width, spermathecal head
round (Fig. 34) to oval (Figs. 32, 36)

DISTRIBUTION. — Southwestern Cameroon and Bioko Island in the Gulf of Guinea (Fig. 63).

MATERIAL EXAMINED. — CAMEROON: Southwest: Fako Division: Mt. Cameroon: mist for-
est on south side, elev. 1425 m, 4°06'28"N, 9°07'10"E, 26-28 January 1992, C. Griswold, J.
Coddington, G. Hormiga, S. Larcher (32 CAS, 2? USNM); grassland near Mann’s Spring, elev.
2050 m, 4°08'N, 9°07'E, 21-25 January 1992, C. Griswold, J. Coddington, G. Hormiga, S. Larcher
(172 CAS, 542 USNM); Buea, type of Mnesitheus zoropsides Strand (1° NHMV); Mapanja,
1884-1885, Knutson and Valdau, type of Mnesitheus asper Thorell (1 ? RMS). Meme Division: Mt.
Kupé above Nyassosso, forest, elev. 800-1200 m, 4°50'N, 9°41'E, 16-19 February 1992, C.
Griswold, N. Scharff, C. Wanzie, S. Larcher, P. Masongo(1 2 CAS). Northwest: Bali, Bafuchu Mbu,
Shum Laka, elev. 1600 m, pitfall, December 1991—February 1992, H. Doutrelepont (1? MRAC),.
EQUATORIAL GUINEA: Bioko: Moca, 3°22'0"N, 8°39'57"E, elev. ca. 1400 m, 6—10 October
1998, K. Dabney and D. Ubick (3°71 2 CAS), 1500 m, “ex: silk-lined burrow on tree fern,” 3—10 Octo-
ber 1998, D. Ubick (1o* CAS); Pico Basilé, 3°37'38"N, 8°48'15"E, elev. ca. 1750 m, in rotting logs,
27-29 September 1998, K. Dabney and D. Ubick (1¢°22 CAS); Moka (sic), types of Mnesitheus
vittatus Simon (3 juveniles, MNHN AR817).

Raecius congoensis, new species
Figures 7; 57, 58, 03

TYPE. — Female holotype from Congo, Lulimbi, embouchure de la riv. Ishasha dans le lac Ed-
ward (Sud-Est) baie de Kyangiro, “dans le sol préléve,” collected July-August 1976, M. Lejeune, de-
posited in MRAC (168.827). Paratype female, same data and deposition except “terreau sous litiére,
galerie forestiere” (MRAC168.862).

ETYMOLOGY. — Refers to occurrence in Congo.

GRISWOLD: AFRICAN RAECIUS SPECIES 125

NOTE. — This species was discussed and illustrated in Griswold (1993:14, figs. 28-30). Data
from the species provided the female Raecius characters in the phylogenetic matrix. This species also
provided exemplar data for Raecius in Griswold et al. (1999).

DIAGNOSIS. — Female with epigynal median lobe weakly concave posteriorly, very broad,
length less than 0.55 times width, lateral lobes without teeth (Fig. 57). Male unknown.

MALE. — Unknown.

FEMALE (Holotype). — Total length 15.73. Carapace orange-brown dorsally, shading to dusky
red-brown laterally and on pars cephalica; sides sparsely covered with dark setae that overlay black
reticulate patterns that radiate from thoracic fovea; thoracic fovea dusky; each eye narrowly sur-
rounded by black pigment; chelicerae dark red-brown except boss orange; labium and pedipalpal
coxae orange-brown with dusky maculations; sternum dusky gray-brown; leg coxae yellow-brown
with dusky markings, these most pronounced on I and II, reticulate basally becoming solid distally;
legs and pedipalpi yellow-brown, legs I and II with dark gray forming longitudinal lateral and dorsal
mottling on femora and tibiae; abdomen yellow-gray, faded, markings appear to have comprised nar-
row dorsal longitudinal light band and light and dark spots laterally. Carapace 7.52 long, 4.38 wide,
2.95 high; clypeus 0.39 high, height 2.05 times AME diameter; ocular area 0.64 long, 2.05 wide, OAL
1.36 times OQL; ratio of eyes AME:ALE:PME:PLE, 1.00:1.18:1.04:1.33, diameter of PME 0.20;
AME-AME interdistance 1.11 times AME diameter, AME-ALE 1.26 times AME; PME-PME
interdistance 1.57 times PME, PME-PLE 2.71 times PME; ALE-PLE equals PLE. Chelicerae 3.10
long. Sternum 3.84 long, 2.04 wide; labium 1.53 long, 1.06 wide; pedipalpal coxae 2.39 long, 0.98
wide. Femur I length 1.10 times carapace width. Spination: typical. ITC I and II absent, III very small,
IV small. Leg measurements (Femur + Patella + Tibia + Metatarsus + Tarsus = [Total]): I: 4.83 + 2.49
ono l 283- 2.34 = [16,10]; 1: 4.05 + 2.204: 2:78 + 2:44 + 2:00= [13.47]; Ml: 3:27 F 1:30 1 Sit
2.15 + 1.56=[10.29]; IV: 4.59 + 2.29 + 3.32 + 3.41 +2.00=[15.61]; pedipalpus: 2.39 + 1.22 + 1.15+
[absent] + 2.07 = [6.83]. Epigynum with median lobe weakly concave posteriorly, very broad, length
less than 0.55 times width, lateral lobes contiguous posteriorly, without teeth (Fig. 57; Griswold 1993,
fig. 28). Vulva with copulatory duct broad with lateral diverticulum, spermathecal head narrow witha
few pores, spermathecal base short, extending mesad to fertilization duct (Fig. 58; Griswold 1993,
figs. 29, 30).

Variation (N=2). Total length 12.53-15.73; ratios of carapace length/width 1.51—1.72,
height/width 0.69—-0.71, PER/OAL 3.22-3.45, OQP/OQA 1.10—1.29; ratios of clypeal height/diame-
ter AME 1.86~—2.05, cheliceral length/clypeal height 7.95—8.33, ratio of length femur I/carapace width
1.09—1.10; number of ventroapical spines on metatarsi I and II ranging from 2 to 5; median and proxi-
mal dorsal femoral spines present or absent; scopulae weak to strong beneath metatarsus II, extending
from 2 to entire length of segment.

DISTRIBUTION. — Southeastern Congo basin (Fig. 63).

MATERIAL EXAMINED. — Only the holotype and paratype.

Raecius crassipes (L. Koch, 1875)
Figure 63

Amaurobius crassipes L. Koch, 1875:32, pl. 3, f. 4 (holotype juvenile from “Ethiopia” “in dem trocken
Strombette des Anseba” [from the dry streambed of the Anseba River, in what is today northern Eritrea], in
BMNH, examined.)

Raecius crassipes, Simon, 1892:230. Roewer 1954:1283. Bonnet 1958:3847. Platnick 2002.

Note. — The juvenile female type specimen is faded, shriveled and disarticulated, but all parts
are present: The relatively long carapace and entire cribellum with linear bunches of cribellar spigots
leave no doubt that it is congeneric with the other species here placed in Raecius. Because the speci-

126 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES
Volume 53, No. 10

men is immature, diagnosis is difficult, but because the type locality is geographically distant and eco-
logically different (i.e., arid) from those of other Raecius species, | consider R. crassipes distinct.

DIAGNOSIS. — The juvenile specimen lacks diagnostic genitalia and is faded. Koch’s figure
(1875, pl. 3, fig. 4) depicts a spider with dark carapace and pale pars cephalica and uniformly pale ab-
domen. Other Raecius species have dark abdomens with at least a faint longitudinal median band. The
4123 leg formula differs from the 1423 of other Raecius.

JUVENILE TYPE FEMALE. — Total length approximately 8.0 mm. Specimen faded, Koch’s figure
(1875, pl. 3, fig. 4) depicts a dark carapace and pale pars cephalica and uniformly pale abdomen. Cara-
pace 4.80 long, 3.15 wide, 1.55 high; clypeus 0.25 high, height 2.27 times AME diameter; ocular area
0.45 long, 1.35 wide, OAL 1.28 times OQL; ratio of eyes AME:ALE:PME:PLE, 1.00:1.72:1.54:1.72,
diameter of PME 0.17; AME-AME interdistance 1.18 times AME diameter, AME-ALE 1.45 times
AME; PME-PME interdistance 0.68 times PME, PME-PLE 1.88 times PME; ALE-PLE 0.26 times
PLE. Chelicerae 2.35 long. Sternum 2.25 long, 1.40 wide, apex forming blunt angle; labium 1.00
long, 0.80 wide; pedipalpal coxae 1.70 long, 0.75 wide. Femur I length 1.06 times carapace width.
Spination typical for female Raecius except femora lacking spines. Scopulae: dense, extending below
apical 1/2 of metatarsi I and II, restricted to tarsi III and IV. ITC present on all legs, ITC HI smallest.
Leg measurements (Femur + Patella + Tibia + Metatarsus + Tarsus = [Total]: I: 3.35 + 1.70 + 2.50 +
1.70+ 1.65=[10.90]; Il: 2.85 + 1.504 2.00+ 1.60+1.60=[9.55]; Il: 2.20+ 140+ 1.25 osamlezs
= [7.75]; IV: 3.25 + 1.65 + 2.65 + 2.50 + 1.60 = [11.65]; pedipalpus: 1.50 + 0.85 + 0.75 + [absent] +
1.40 = [4.50].

DISTRIBUTION. — Known only from the type locality, in an arid region of northern Eritrea (Fig.
63).

MATERIAL EXAMINED. — Only the type.

Raecius jocquei, new species
Figures 10-17, 19, 21-23, 43-53, 63

TyPE. — Male holotype from Céte d’Ivoire, Appouesso, forét classée de la Bossematie, pitfall
traps, 31 October 1994, R. Jocqué and N. Seabé, deposited in MRAC (202.373). Paratypes, same lo-
cality, 18°42 MRAC, 11? CAS.

ETYMOLOGY. — Named in honor of Rudy Jocqué, who collected these specimens as well as
many other rare and interesting African Arachnida.

NoTE. — This species provided partial exemplar data for Raecius in Griswold et al. (1999).

DIAGNOSIS. — Male with pedipalpal bulb (Figs. 45-47, 50-53) having median apophysis ex-
panded and bilobate at apex and STP prominent, curved and narrow, pedipalpal tibia with RTA large,
expanded and bilobate at apex, greater than 0.85 times the length of the tibia. Female with epigynal
median lobe deeply concave posteriorly, U-shaped, length 0.80—0.85 times width (Fig. 48).

MALE (Holotype). — Total length 6.76. Spider yellow-gray, unmarked except carapace dusky
laterally overlain sparsely with dark setae, leaving median pale band, broad on pars cephalica and nar-
row on pars thoracica; each eye surrounded with black pigment; dorsal process of pedipalpal tibia
dark brown; dorsum of cymbium dusky; abdomen dark gray dorsally except for narrow median pale
band, sides mottled. Carapace 4.00 long, 2.63 wide, 1.27 high; clypeus 0.18 high, height 1.13 times
AME diameter; ocular area 0.40 long, 0.97 wide, OAL 1.21 times OQL; ratio of eyes
AME:ALE:PME:PLE, 1.10:1.30:1.20:1.00, diameter of PME 0.17; AME interdistance 0.45 times
AME diameter, AME-ALE 0.27 times AME; PME interdistance 0.33 times PME, PME-PLE 1.08
times PME; ALE-PLE 0.50 times PLE. Chelicerae 1.39 long. Sternum 1.98 long, 1.35 wide; labium
0.73 long, 0.55 wide; pedipalpal coxae 1.25 long, 0.42 wide. Femur I length 1.33 times carapace
width. Spination: basic pattern except femora I-IV d1-1-1, tibia I dl-1-1, metatarsi IIT and IV p1-1-2.
ITC I small, slender, II small, stout, III and IV large. Leg measurements (Femur + Patella + Tibia +

GRISWOLD: AFRICAN RAECIUS SPECIES 127

Metatarsus + Tarsus = [Total]): I: 3.49 + 1.49 + 3.76 + 3.45 + 1.96 =[14.15]; Il: 2.86 + 1.37 +2.59 +
peso 73 — [1.14]; WI: 2.31 4+ 1.144 1.49 +2.08 + 1.43 =[8.45];1V23.29 +145 22.9453 10:
1.73 = [12.51]; pedipalpus: 1.49 + 0.71 + 0.73 + [absent] + 1.14 = [4.07]. Pedipalpus (Figs. 45-47,
50-53) with tibia with RTA large, expanded and bilobate at apex, length greater than 0.85 times that of
tibia, VTA conical, broad and blunt, pro- and retroventral margins of tibia at tibia-tarsus joint pro-
duced into small processes; bulb with median apophysis expanded and bilobate at apex, with 2 STP,
one (STP1) prominent, curved and narrow, extending nearly to apex of bulb, a smaller broad, curved
process (STP2) arising behind STP1, embolus slender, ribbon-like, conductor short.

Variation (N=5). Total length 5.36—-6.76; ratios of carapace length/width 1.49—-1.62,
height/width 0.38—0.48, ratios of PER/OQP 2.39-2.65, PER/OAL 2.34—2.53, OQP/OQA 1.07-1.25,
ratios of clypeal height/diameter AME 0.84—1.78, cheliceral length/clypeal height 7.3 1—8.36, ratio of
length femur I/carapace width 1.32—2.84. Markings may be bold to faint. Retrodorsal spines on
femora I-III present or absent.

FEMALE (Paratype). — Total length 9.23. Carapace red-brown, sides dusky with black reticulate
markings, these markings conspicuous on sides of pars cephalica and behind posterior eyes;
chelicerae red-brown with longitudinal black striae, boss orange; labium and pedipalpal coxae or-
ange-brown, paler at bases and tips; sternum dusky yellow-gray; leg coxae gray except reticulated
with yellow-gray basally; legs and pedipalpi red-brown, leg femora dusky laterally; abdomen dark
gray on dorsum and sides except for narrow central longitudinal white band, venter and spinnerets
pale yellow-gray. Carapace 5.05 long, 2.95 wide, 1.71 high; clypeus 0.31 high, height 1.72 times
AME diameter; ocular area 0.40 long, 1.40 wide, OAL 1.21 times OQL; ratio of eyes
AME:ALE:PME:PLE, 1.00:1.07:1.00:1.15, diameter of PME 0.18; AME-AME interdistance 0.72
times AME diameter, AME-ALE 0.77 times AME; PME-PME interdistance 0.54 times PME,
PME-PLE 1.69 times PME; ALE-PLE 0.33 times PLE. Chelicerae 1.98 long. Sternum 2.36 long, 1.62
wide; labium 1.03 long, 0.77 wide; pedipalpal coxae 1.64 long, 0.61 wide. Femur I length 1.18 times
carapace width. Spination: typical. ITC I small, II-IV large. Leg measurements (Femur + Patella +
Tibia + Metatarsus + Tarsus = [Total]): I: 3.49 + 1.65 + 2.78 + 2.39 + 1.92 = [12.23]; II: 3.14+ 1.61 +
Pee 206 121.59 = [10571 2M 2.434 1129 +129 + 1.84 + 1.14 =[7.99]; IV: 3.45:-+ 1612754
2.59 + 1.37 =[11.77]; pedipalpus: 1.65 + 0.82 + 0.82 + [absent] + 1.37 = [4.66]. Epigynum with me-
dian lobe deeply concave posteriorly, U-shaped, lateral lobes separate posteriorly, with
mesad-extending teeth near level of anterior margin of median lobe (Fig. 48). Vulva with copulatory
duct narrow, simple, spermathecal head narrow with several pores, spermathecal base very short, ex-
tending basad to fertilization duct (Fig. 49).

Variation (N=4). Total length 4.78-9.23; ratios of carapace length/width 1.59-1.77,
height/width 0.47—-0.64, ratios of PER/OQP 2.51—3.13, PER/OAL 3.09-3.50, OQP/OQA 0.93-1.17,
ratios of clypeal height/diameter AME 1.58—1.72, cheliceral length/clypeal height 6.3 1—7.67, ratio of
length femur I/carapace width 1.08—1.87. Markings faint to bold. Epigynal teeth small and limited to
LL or extending mesad over margins of ML, ML length 0.80—0.85 times width.

DISTRIBUTION. — Cote d’ Ivoire (Fig. 63).

MATERIAL EXAMINED. — COTE D’ IVOIRE: Appouesso, forét classée de la Bossematie, pit-
fall traps, R. Jocqué and N. Séabé, 3 May 1994 (8c°1 2 paratypes MRAC), 18 May 1994 (paratypes:
472? MRAC, 1°12? CAS), 31 October 1994 (2¢ paratypes, 1% holotype MRAC), 1 December 1994
(1% paratype MRAC), 18 March 1994 (1c paratype MRAC); Abengourou, forét classée de la
Bossematie, primary forest, pi¢ge, 4 March and 18 May 1994, N. Séabé (2° MRAC).

128 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES
Volume 53, No. 10

Raecius scharffi, new species
Figures 4—5, 9, 54-56, 59-63

TYPES. — Male holotype from pitfall traps at 750 m elevation in lowland rain forest in Uzungwa
Scarp Forest Reserve above Chita Village, Iringa Region, Tanzania, collected on 25—29 October 1984
by Nikolaj Scharff, deposited in ZMUC. Paratype males: two from Kambai Forest Reserve and two
from Marimba Forest Reserve, Tanzania, one from Marimba in CAS, and remainder in ZMUC.

ETYMOLOGY. — Named in honor of Nikolaj Scharff, who collected these specimens as well as
many other rare and interesting African Arachnida.

DIAGNOsIS. — Male with pedipalpal bulb having median apophysis produced into distal hook
and STP reduced, pedipalpal tibia with DTA, RTA small, attenuate at apex, less than 0.4 times the
length of the tibia (Figs. 54—56, 59-62). Female unknown.

MALE (Holotype). — Total length 7.33. Spider pale yellow-gray, unmarked except dark brown
at lateral margins of carapace above chelicerae and apices of chelicerae, carapace sparsely covered
with dark setae, these becoming densest laterally; abdomen gray-brown on dorsum and sides, with
faint anteromedian light band, venter and spinnerets yellow-white. Carapace 4.09 long, 2.76 wide,
1.53 high; clypeus 0.18 high, height 1.29 times AME diameter; ocular area 0.31 long, 1.06 wide, OAL
1.11 times OQL; ratio of eyes AME:ALE:PME:PLE, 1.00:1.11:1.11:1.11, diameter of PME 0.16;
AME interdistance 0.50 times AME diameter, AME-ALE 0.70 times AME; PME interdistance 0.64
times PME, PME-PLE 1.36 times PME; ALE-PLE 0.27 times PLE. Chelicerae 1.76 long. Sternum
1.96 long, 1.33 wide; labium 0.86 long, 0.55 wide; pedipalpal coxae 1.33 long, 0.47 wide. Femur I
length 1.71 times carapace width. Spination: basic pattern except pedipalpal patella has p0-1-0, tibia
and metatarsus I lack retrolateral spines, tibia II v1-1-2, metatarsus II p0-1-2, metatarsus IV v2-2-1.
ITC I and II very small, slender, HI and IV small. Leg measurements (Femur + Patella + Tibia +
Metatarsus + Tarsus = [Total]): I: 4.71 + 1.61 + 4.77 + 3.94 + 2.65 = [17.68]; II: 3.87 + 1.42 + 3.42 +
3.10+2.06=[13.87]; IL: 3.23 + 1.23 + 2.13 +2.65 + 1.55 =[10.79]; IV: 4.52 + 1.49 + missing + miss-
ing + missing = [not calculated]; pedipalpus: 1.68 + 0.71 + 0.58 + [absent] + 1.29 = [4.26]. Pedipalpus
(Figs. 54—56, 59-62) with tibia with a small, conical DTA, RTA short, curved, bluntly pointed, length
less than 0.4 times that of tibia, VTA small and low, and retroventral margin of tibia at tibia-tarsus
joint produced into small process; bulb with median apophysis produced into distal hook, STP re-
duced, represented only by a membranous swelling, embolus short, and conductor large.

Variation (N = 5). Total length 6.67—8.40; ratios of carapace length/width 1.48—1.90,
height/width 0.52—0.74, ratios of PER/OQP 2.46—2.92, PER/OAL 3.34—3.71, OQP/OQA 1.02-1.27,
ratios of clypeal height/diameter AME 1.13—1.29, cheliceral length/clypeal height 9.78—12.00, ratio
of length femur I/carapace width 1.65—2.17, leg formulae of paratype males 1423. Paratype males
from Marimba and Kambai have the abdomen slightly darker than the type: yellow-gray dorsally with
anteromedian light longitudinal band and faint transverse gray bands posteriorly. Spination:
distodorsal spines on femora HI and 1V and metatarsus IV present or absent. DTA of pedipalpal tibia
small and conical (holotype) or long and digitate (paratypes).

FEMALE. — Unknown.

DISTRIBUTION. — Eastern Arc mountains of Tanzania (Fig. 63).

MATERIAL EXAMINED. — TANZANIA: Iringa Region: Uzungwa Scarp Forest Reserve above
Chita Village, elev. 750 m, pitfall traps in lowland Rain Forest, 25—29 October 1984, N. Scharff (1
holotype * ZMUC). Tanga Region: Muheza District, Kambai Forest Reserve, 4°59’S, 38°41’E, pit-
fall traps, Frontier Tanzania Expedition (2 paratype &* ZMUC), Marimba Forest Reserve, 5°01'S,
38°41'E, pitfall traps, Frontier Tanzania Expedition (2 paratype «, | ZMUC and | CAS); East
Usambara Mountains, 12km SE Amani, Kihuhwi-Zigi Forest Reserve, 5°6.3'S, 38°40.6'E,
400-450 m elev., 2-4 November 1995, C. Griswold (1 juvenile, CAS).

GRISWOLD: AFRICAN RAECIUS SPECIES 129

EXCLUDED SPECIES
Raecius aculeatus Dahl, 1901

Raecius aculeatus Dahl, 1901a:188 (holotype juvenile from between Knako and Kimpoko, Congo, labeled
Raecius spinosus, in ZMB, examined). Roewer 1954:1283. Bonnet 1958:3846. Platnick 2002.
Raecius spinosus Blumenthal, 1935:709.

Note. — No specimen has been located labeled as Raecius aculeatus but there is a specimen in
the Zoologisches Institut der Humboldt Universitat, Berlin, labeled ““Raecius spinosus F. Dahl,
Kimpoko, Biittner, ZMB25278.” Blumenthal (1935:709) referred to this specimen of Raecius
spinosus and Bonnet (1958:3846) noted that Blumenthal’s work refers to the specimen described by
Dahl as R. aculeatus. The specimen is a juvenile just prior to molting (no mature structures are visible
through the cuticle) and is disarticulated. Part was formerly dry and on a pin, but has been returned to
alcohol and other parts are slide mounted. The parts in alcohol include an intact carapace, left legs
LIV and a partial, shriveled abdomen. The slide-mounted parts comprise the chelicerae, labium and
pedipalpi, right legs and spinning organs. The calamistrum is oval, cribellar spigots are in linear
groups and the eyes are in two, nearly straight rows, suggesting placement in Zorocratidae. The cara-
pace is long and narrow (length 1.8 times width) suggesting Raecius, but the cribellum is divided (not
entire) and feathery hairs are lacking. Because this specimen does not share these synapomorphies
with Raecius, | suggest that it should be excluded from Raecius and be placed incertae sedis in
Zorocratidae.

ACKNOWLEDGMENTS

Principal support for this project came from National Science Foundation grants BSR-9020439
and DEB-9020439. The contributions of Robin Carlson (silk fine structure and spinning behavior)
and Hillary Guttman (specimen measuring and mapping) were made possible by the California Acad-
emy of Sciences (CAS) Summer Systematics Institute, supported by NSF grant BIR-9531307. Re-
search in Equatorial Guinea (Bioko Island) was made possible by a grant from Central African
Regional Programs for the Environment (CARPE) to Robert Drewes and C. Griswold. Additional
support came from the Exline-Frizzell and In-house Research Funds of CAS and the Schlinger Foun-
dation.

Illustrations are by Jenny Speckels (JS) or Virginia Kirsch (GK) and are attributed in the figure
captions. Gustavo Hormiga kindly supplied photos of the living spiders. Assistance with manuscript
preparation was provided by Joel Ledford, Heather Tennison, and Darrell Ubick. A draft of the manu-
script was read and criticized by Diana Silva, Darrell Ubick, and two anonymous reviewers.

Specimens and observations essential to the completion of this study were obtained through
fieldwork in Cameroon, Equatorial Guinea and Tanzania. This fieldwork was made possible through
permissions granted by a variety of government bodies and traditional authorities. Permits to do re-
search in Cameroon were granted by the Institute of Zootechnical Research, Ministry of Higher Edu-
cation and Scientific Research of the Republic of Cameroon. | especially thank Dr. John T. Banser,
Director of the Institute, and Dr. Chris Wanzie, for facilitating my research in Cameroon. Fieldwork in
Cameroon was also made possible by the permission and assistance from the following traditional au-
thorities: Their Royal Highnesses the chiefs of Batoke, Etome, and Mapanja, H.R.H. R. M. Ntoko,
Paramount Chief of the Bakossi, Chiefs Jonas Achang, Albert Ekinde, and Peter Epie of Nyassosso.
Research in Tanzania was made possible through a Research Permit from the Tanzania Commission
for Science and Technology (COSTECH) and Residence Permit Class C from the Tanzanian Depart-
ment of Immigration, and export of specimens made possible by a CITES Exemption Certificate from

130 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES
Volume 53, No. 10

the Wildlife Division of the United Republic of Tanzania, facilitated by Professor Kim M. Howell of

the University of Dar-es-Salaam. Research in the West Usambaras (Tanzania) was made possible by

Dr. S. A. O. Chamshama, Dean of Forestry, Sokoine University, and Mr. Modest S. Mrecha, Officer

in Charge, Mazumbai Forest Reserve. I thank Robert Drewes, Jens Vindum, and especially D. Keith

Dabney and Darrell Ubick for carrying out fieldwork on Bioko Island.
The following individuals and institutions are gratefully acknowledged for loan of material:

Janet Beccaloni, Arachnida and Myriapoda Section, Department of Zoology, The Natural History
Museum, London, England (BMNH)

Jonathan Coddington, National Museum of Natural History, Smithsonian Institution, Washington
DC, USA (USNM)

Jurgen Gruber, Naturhistorisches Museum, Zoologisches Abteilung, Vienna, Austria (NHMV)

TorbjOrn Kronestedt, Riksmuseets Entomologiska, Stockholm, Sweden (RMS)

Rudy Jocqué, Musée Royal de L’ Afrique Centrale, Tervuren, Belgium (MRAC)

Jason Dunlop, Museum fiir Naturkunde, Zoologisches Institut der Humboldt Universitat, Berlin, Ger-
many (ZMB)

Gisella Rack, Zoolgisches Institut und Zoologisches Museum, Universitat Hamburg, Hamburg, Ger-
many (ZMH)

Christine Rollard and the late Jacqueline Heurtault, Muséum National d’Histoire Naturelle,
Laboratoire des Arthropodes, Paris, France (MNHN)

Nikolaj Scharff, Zoologisk Museum, Copenhagen, Denmark (ZMUC)

Other material is in the collection of the California Academy of Sciences, San Francisco (CAS).
This is contribution 22 from the California Academy of Sciences Center for Biodiversity Research
and Information (CBRI).

GRISWOLD: AFRICAN RAECIUS SPECIES 131

FIGURE 1. Raecius asper, female from Mann’s Spring, habitus, dorsal. Illustration by JS. Scale bar = 1.0 mm.

PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES
Volume 53, No. 10

FIGURES 2-3.

Raecius asper, female from Mann’s Spring. 2. Lateral. 3. Ventral. Illustrations by JS. Scale bar = 1.0 mm.

GRISWOLD: AFRICAN RAECIUS SPECIES 133

FicureEs 4-9. Legs of Raecius. 4, 5,9. R. scharffi, male holotype. 6, 8. R. asper, female from Mann’s Spring. 7. R. congoensis,
holotype. 4. Trichobothrial base, tibia IV. 5. Tarsal organ IV. 6. Close up of calamistral setae (center), with plumose and feath-
ery setae at right. 7. Tarsus I, showing tarsal organ (line), trichobothrial base, and plumose setae. 8. Metatarsus IV showing
calamistrum. 9. Tibia IV showing plumose and feathery setae. Scale bars: 4-5 = 10 um, 6-7 = 50 pm, 8= 1.0 mm, 9= 100 ym.

134 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES
Volume 53, No. 10

Ficures 10-13. Right spinnerets of female Raecius jocquei paratype. 10. Overview. 11. ALS. 12. PMS. 13. PLS. AC =
aciniform gland spigot(s), CY = cylindrical gland spigot(s), MAP = major ampullate gland spigot(s), mAP = minor ampullate
gland spigot(s), PI = piriform gland spigot(s). Scale bars: 10 = 150 wm, 11-13 = 43 pm.

GRISWOLD: AFRICAN RAECIUS SPECIES 135

Figures 14-17. Left spinnerets of male Raecius jocquei paratype. 14. Overview. 15. ALS. 16. PMS. 17. PLS. AC =
aciniform gland spigot(s), MAP = major ampullate gland spigot(s), mAP = minor ampullate gland spigot(s), N = nubbin of
pseudoflagelliform gland spigot(?), PI = piriform gland spigot(s). Scale bars: 14 = 150 um, 15-17 = 43 pm.

136 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES
Volume 53, No. 10

LPBZAI SERS SADN

S = gee

SIN ‘ i |

J \
OP? 0 / e GE. ID \ OS See

FIGURES 18-21. Raecius female spinning organs. 18, 20. R. asper, Mann’s Spring, Cameroon. 19, 21. R. jocquei paratype.
18. Spinnerets and cribellum, overview. 19. Cribellum. 20. Left PLS. 21. Close of up groups of cribellar spigots. AC =
aciniform gland spigot(s), CY = cylindrical gland spigot(s), PsFl = pseudoflagelliform gland spigot. Scale bars: 18 = 500 um,
19 =201 pm, 20 = 100 pm, 21 = 20 pm.

GRISWOLD: AFRICAN RAECIUS SPECIES 137

FX STP ZEN

FiGuRES 22-23. Expanded right pedipalpus of male Raecius jocquei paratype. 22. Retrolateral. 23. Prolateral. AN = anelli, C
= conductor, E= embolus, MA = median apophysis, P = petiole, RTA = retrolateral apophysis of tibia, ST = subtegulum, STP =
sclerotized tegular process, T=tegulum, VTA = ventral apophysis of tibia. Straight white lines on Fig. 23 to locking lobes of ST
and T. Illustrations by JS. Scale bar = 0.5 mm.

‘PIOMSLID sapteyD Aq 9Z ‘eSIULIOH] OARISND Aq /Z pure CZ ‘PZ
SOINBI{ (PUNO UO UOWTUOS 919M SMO.LING SNiJavY “WU YO [Z Ie PuL]sseis pouing */Z ‘Sdo] SuN}04 ut punoy ATPLoIdAy smog sn12avy “WI CYP] 1 ISOOJ IST ‘OZ “WOOT Z ‘pues
“HTES O7B][IQL JO Ie]JOS YM MOLING “CZ “WOOT “Suudg s uueypy saoqe woy ‘ajeway “pZ ‘WOOIDWIeD “YAY Woy syeyIqey pure sqam ‘suoundads :wadsy snizavy *LZ7—pZ STU!

Volume 53, No. 10

nN
eal
0
Z
S
'S)
nA
Ly,
S
4
=.
aa)
A
<
O
<
Ss
Zz
~
©
J
<
iS)
aa]
Ly
[=
Ly
S)
nN
Oo
Ez
Q
aa
aa
v
e)
(a4
AL

GRISWOLD: AFRICAN RAECIUS SPECIES 139

FIGURES 28-32. Raecius asper. 28-30. Male from Moca, Bioko Island, left pedipalpus. 28. Prolateral. 29. Retrolateral. 30.
Ventral. 31-32. Female from Mann’s Spring, Mt. Cameroon. 31. Epigynum, ventral. 32. Vulva, dorsal. Illustrations 28-30 by
GK, others by JS. Scale bars = 0.25 mm.

140 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES
Volume 53, No. 10

FIGURES 33-36. Raecius asper, female genitalia. 33-34. Bafuchu Mbu, Cameroon. 35-36. Mt. Cameroon, mist forest at
1400m. 33, 35. Epigyna, ventral. 34, 36. Vulvae, dorsal. Illustrations by JS. Scale bar = 0.25 mm.

GRISWOLD: AFRICAN RAECIUS SPECIES 141

a 77
meu |

~dQ._

FIGURES 37-40. Raecius asper male from Moca, Bioko Island, left pedipalpus. 37. Prolateral. 38. Retrolateral. 39. Ventral.
40. Apicoventral. C = conductor, E = embolus, MA = median apophysis, RTA = retrolateral apophysis of tibia, STP =
sclerotized tegular process, VTA = ventral apophysis of tibia. Scale bars 37-39 = 600 um, 40 = 300 um.

142 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES
Volume 53, No. 10

eee

NGI C5
=

“ee

=f "at
Nak he
— na Pa \oumeaw i 7. a
FIGURES 41—44. Raecius spp., abdomens. 41—42. R. asper female from Mann’s Spring, Mt. Cameroon. 43-44. R. jocquei,
paratypes. 41. Epigynum, ventral. 42. Vulva, dorsal. 43. Venter of 2 abdomen, showing plumose and feathery hairs. 44. Male

epiandrous region. CD = copulatory duct, EF = epigynal fold, FD = fertilization duct, HS = head of spermatheca, LL = lateral
lobes of epigynum, ML = median lobe of epigynum. Scale bars: 41 = 300 um, 42 = 250 um, 43 = 30um, 44 = 75 um.

GRISWOLD: AFRICAN RAECIUS SPECIES 143

FIGURES 45-49. Raecius jocquei, Abengourou. 45-47. Left male pedipalpus. 45. Prolateral. 46. Retrolateral. 47. Ventral. 48.
Epigynum, ventral. 49. Vulva, dorsal. Illustrations by JS. Scale bars = 0.25 mm.

144 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES
Volume 53, No. 10

rm SSS

FiGureEs 50-53. Raecius jocquei, Abengourou, right male pedipalpus. 50. Retrolateral. 51. Prolateral. 52. Ventral. 53. Dor-
sal. C = conductor, E = embolus, MA = median apophysis, RTA = retrolateral apophysis of tibia, STP (1 and 2) = sclerotized
tegular processes | and 2, VTA = ventral apophysis of tibia. Scale bars: 50, 53 = 430 tm; 51, 52 = 300 um.

GRISWOLD: AFRICAN RAECIUS SPECIES 145

FiGuRES 54-58. Raecius spp. 54—5S6. R. scharffi, holotype, left male pedipalpus. 54. Prolateral. 55. Retrolateral. 56. Ventral.
57-58. R. congoensis, female holotype. 57. Epigynum, ventral. 58. Vulva, dorsal. DTA = dorsal apophysis of tibia. Illustrations
by JS. Scale bars = 0.25 mm.

146 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES
Volume 53, No. 10

FiGures 59-62. Raecius scharffi, holotype, right male pedipalpus. 59. Retrolateral. 60. Prolateral. 61. Ventral. 62.
Apicoventral. C = conductor, E = embolus, MA = median apophysis, RTA = retrolateral apophysis of tibia, VTA = ventral
apophysis of tibia. Scale bars: 59 = 600 um; 60, 61 = 430 um; 62 = 200m.

GRISWOLD: AFRICAN RAECIUS SPECIES 147

Mi congoensis
@ crassipes
A jocquei
W. scharfti

‘
1
!

FIGURE 63. Map of Africa, showing locality records for Raecius species.

148 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES
Volume 53, No. 10

LITERATURE CITED

ASHER, R. J. 1997. African and Malagasy tenrecs: a biogeographic parallel with lemuriform primates? American
Journal of Physical Anthropology, supplement 24:69—70.

BLUMENTHAL, H. 1935. Untersuchungen tiber das “Tarsalorgan” der Spinnen. Zeitschrift ftir Morphologie und
Okologie der Tiere 29(5):667—719.

BONNET, P. 1957. Bibliographia Araneorum. Toulouse 2 (3):1927—3026.

. 1958. Bibliographia Araneorum. Toulouse 2 (4):3027—4230.

BRIGNOLI, P. M. 1983. A catalogue of the Araneae described between 1940-1981. Manchester University Press,
Manchester. 755 pp.

DAHL, F. 1901a. Uber den Werth des Cribellums und Calamistrums fiir das System der Spinnen und eine
Uebersicht der Zoropsiden. Sitzungsberichte der Gesellschaft naturforschender Freunde zu Berlin
1901:177-199.

. 1901b. Nachtrag zur Uebersicht der Zoropsiden. Sitzungsberichte der Gesellschaft naturforschender

Freunde zu Berlin 1901:244—255.

.1913. Vergleichende Physiologie und Morphologie der Spinnentiere unter besonderer
Beriicksichtigung der Lebensweise I. 113 pp. Jena.

EBERHARD, W. G. 1988. Combing and sticky silk attachment behavior by cribellate spiders and its taxonomic im-
plications. Bulletin of the British Arachnological Society 7(8):247—251.

GRISWOLD, C.E. 1985. A revision of the African spiders of the family Microstigmatidae (Araneae:
Mygalomorphae). Annals of the Natal Museum 27:1—37.

. 1987a. A revision of the jumping spider genus Habronattus F. O. P.-Cambridge (Araneae; Salticidae),

with phenetic and cladistic analyses. University of California Publications in Entomology 107:1—344.

. 1987b. The African members of the trap-door spider family Migidae (Araneae: Mygalomorphae). 1:

The genus Moggridgea O. P.- Cambridge, 1875. Annals of the Natal Museum 28 (1):1—118.

. 1987c. The African members of the trap-door spider family Migidae (Araneae: Mygalomorphae). 2: the

genus Poecilomigas Simon, 1903. Annals of the Natal Museum 28:475—497.

. 1987d. A review of the southern African spiders of the family Cyatholipidae Simon, 1894 (Araneae:

Araneomorphae). Annals of the Natal Museum 28:499-542.

. 1990. A revision and phylogenetic analysis of the spider subfamily Phyxelidinae (Araneae,

Amaurobiidae). Bulletin of the American Museum of Natural History 196:1—206.

. 1991a. Cladistic Biogeography of Afromontane Spiders. Australian Systematic Botany 4 (1):73-89.

.1991b. A revision and phylogenetic analysis of the African spider genus Machadonia Lehtinen

(Araneae: Lycosoidea). Entomologica Scandinavica 22:305—351.

. 1993. Investigations into the phylogeny of the lycosoid spiders and their kin (Arachnida: Araneae:

Lycosoidea). Smithsonian Contributions to Zoology 539:1—39.

. 1994. A revision and phylogenetic analysis of the African spider genus Phanotea Simon (Araneae:

Lycosoidea). Annales, Musée Royal de |’ Afrique centrale, Sciences Zoologiques 273:1—83.

. 1997a. The spider family Cyatholipidae in Madagascar (Araneae, Araneoidea). Journal of Arachnology

25:53-83.

.1997b. Scharffia, a remarkable new genus of spiders from East African Mountains (Araneae,
Cyatholipidae). Journal of Arachnology 25:269-287.

——.1998a. The nest and male of the trap-door spider Poecilomigas basilleupi Benoit, 1962 (Araneae,
Migidae). Journal of Arachnology 26:142—148.

———.1998b. Wanzia fako, a new genus and species of spider from Cameroon (Araneae, Cyatholipidae).
Entomologica Scandinavica 29:121—130.

. 2000. Afromontane spiders in Madagascar (Araneae, Araneomorphae: Cyatholipidae, Phyxelididae,

Zorocratidae). Pp. 345-354 in Diversity and Endemism in Madagascar, W. R. Lourengo and S. M. Good-

man, eds. Mémoires de la Société de Biogéographie, Paris.

. 2001. The living world genera and Afrotropical species of cyatholipid spiders (Araneae, Orbiculariae,
Cyatholipidae). Memoirs of the California Academy of Sciences, Number 26. 251 pp.

GRISWOLD, C. E., J. A. CODDINGTON, N. I. PLATNICK, AND R. R. FORSTER. 1999. Towards a phylogeny of
entelegyne spiders (Araneae, Araneomorphae, Entelegynae). Journal of Arachnology, 27:53—63.

GRISWOLD: AFRICAN RAECIUS SPECIES 149

GRISWOLD, C. E. AND J. LEDFORD. 2001. A monograph of the migid trap-door spiders of Madagascar, with a phy-
logeny of world genera (Araneae, Mygalomorphae, Migidae). Occasional Papers of the California Acad-
emy of Sciences 151:1—120.

GRISWOLD, C. E. AND N. I. PLATNICK. 1987. On the first African spiders of the family Orsolobidae (Araneae,
Dysderoidea). American Museum Novitates, 2892:1—14.

GRISWOLD, C. E. AND D. UBICK. 1999. Spiders—Araneae. Pp. 7-67 in Arthropod surveys on Bioko, Equatorial
Guinea. Report to Central African Regional Programs for the Environment. Online publication at:
http://www.calacademy.org/research/entomology/Entomology%20Resources/entomology_re-
sources.htm.

Kocu, L. 1875. Aegyptische und abyssinische Arachniden gesammelt von Herrn C. Jickeli. Nurnberg, pp. 1-96.

LEHTINEN, P. T. 1967. Classification of the cribellate spiders and some allied families, with notes on the evolu-
tion of the suborder Araneomorpha. Annales Zoologici Fennici, 4:199-468

_ 1975. Notes on the phylogenetic classification of Araneae. Proceedings of the 6th international
arachnological Congress, Amsterdam. 1974:26-29.

MCCALL, R. A. 1997. Implications of recent geological investigations of the Mozambique Channel for the mam-
malian colonization of Madagascar. Proceedings of the Royal Society of London, 264:663—665.

PETRUNKEVITCH, A. 1928. Systema Aranearum. Transactions of the Connecticut Acadademy of Arts, Sciences
and Letters, 29:1—270.

PLATNICK, N. I. 1989. Advances in spider taxonomy: a supplement to Brignoli’s A Catalogue of the Araneae de-
scribed between 1940 and 1981. Manchester University Press, Manchester. 673 pp.

. 1993. Advances in spider taxonomy, 1988-1991: with synonymies and transfers 1940-1980. New York

Entomological Society, New York. 846 pp.

. 1997. Advances in spider taxonomy, 1992-1995: with redescriptions 1940-1980. New York Entomo-

logical Society, New York. 976 pp.

. 2002. The world spider catalog, version 2.5. American Museum of Natural History, online at
http://research.amnh.org/entomology/spiders/catalog8 1-87/index.html.

ROEWER, C. F. 1954. Katalog der Araneae von 1758 bis 1940, Vol. 2a, pages 1-923. Bruxelles.

SIERWALD, P. 1989. Morphology and ontogeny of female copulatory organs in American Pisauridae, with special
reference to homologous features (Arachnida: Araneae). Smithsonian Contributions to Zoology, 484: 1—24.

SIMON, E. 1892. Histoire Naturelle des Araignées, 1(1), Roret, Paris, pp. 1-256.

. 1903. Histoire Naturelle des Araignées, 2(4), Roret, Paris, pp. 669-1080.

. 1907. Arachnides recueillis par L. Fea sur la Céte occidentale d’ Afrique. Ire partie. Annali del Museo
Civico di Storia Naturale di Genova, (3) 3:218—323.

STRAND, E. 1916. Uber einige Arachniden aus Buea in Kamerun. Schaftliche Ergebnisse der Deutschen Zentral
Afrika Expedition 1907-1908, unter Fiihrung Adolg Friedrichs, Herzogs zu Mecklenberg. Archiv fur
Naturgeschichte, 81(A11):139-149.

THORELL, T. 1899. Araneae Camerunenses (Africae occidentalis) quas anno 1891 collegerunt Cel. Dr Y. Sjostedt
aliique. Bihang Till Kongelige Svenska Vetenskaps-Akademiens Handlingar, 25(1):1—105.

Wuite, F. 1978. The afromontane region. Pp. 464-513 in Biogeography and Ecology of Southern Africa,
M. J. A. Werger, ed. Junk, The Hague.

. 1983. The vegetation of Africa: a descriptive memoir to accompany the Unesco/AETFAT/UNSO vege-
tation map of Africa. United Nations Educational, Scientific and Cultural Organization, Paris. 356 pp.

Yooper, A., M. CARTMILL, M. RUVOLO, K. SMITH, AND R. VILGALYS. 1996. Ancient single origin for Malagasy
primates. Proceedings of the National Academy of Sciences, USA, 93:5122—5126.

Ate Fria EPISeeAR

“>

chap AGT waar GALANT lees cedtign vite St beop a
(aurnp, grat hg wil] De gee] boutiet ict) dettagite

eat diate , at a hos eel ay ors
: Ss ae eee cep NIA bn
Rare he Fi Pay ra Bry hy. sagen ie i To oy” whinge ne ay -
¥ a ei Late DP as wi : pi) nie ay orl pee PA en ‘i
" ® Oh: ii (lis 2) 7 at) Bw At i Pare t

7 > ' - y a7 he
a , gegen hol A 1 uy aia a ; 9 hrs |
2 o\m « ov sae on on Pu _ i.» a oe a) a
- ot Prat
i <a [? ; i 7 ‘teil ww
oad | Riget
an 7 , nl be is _ } Perit
Q — 7
hs
AS pal :

a’

PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES

Volume 53, No. 11, pp. 151—160, 2 figs. August 22, 2002

First records of snailfishes (Pisces: Liparidae)
from the Galapagos Islands, with descriptions of two new species,
Paraliparis darwini and Paraliparis galapagosensis

by
David L. Stein
NOAA/OAR R/OSS3
1315 East West Highway, Rm. 11805
Silver Spring, Maryland 20910
email: david.stein@noaa.gov

and

Natalia V. Chernova
Zoological Institute
Russian Academy of Sciences
199034 St. Petersburg, Russia
email: antarct(@zin.ru

Snailfishes (Family Liparidae, Order Scorpaeniformes) are very widely distributed both
in depth and geographically; more than 130 species are known from the Southern Hemi-
sphere, and five species from deep, eastern Pacific, equatorial waters. We report herein the
first record of the family from the Galapagos Islands, and describe two new species,
Paraliparis darwini and P. galapagosensis.

Liparid fishes are distributed worldwide in cold and cool waters at depths from the intertidal to
over 7000 m. They are highly speciose and morphologically diverse, occupying a wide variety of hab-
itats including benthic, epibenthic, and pelagic zones. In the Southern Hemisphere, there are at least
130 described species (Andriashev 1986; Stein and Tompkins 1989; Stein and Andriashev 1990;
Stein, Melendez, and Kong 1991; Andriashev and Stein 1998; Stein, Chernova, and Andriashev 2001;
Chernova and Stein in press; and others) from the Antarctic, Australia, Chile, Argentina, and other lo-
cations. In the tropical and subtropical Pacific, the five known species are from great depths
(Careproctus longifilis Garman 1892, SW of Panama, 3343 m; Paraliparis fimbriatus Garman 1892,
SW of Panama, 3241 m; P. Jatifrons Garman 1899, Panama Bay, 3279 m; P. angustifrons Garman
1899, Panama Bay, 935 m; P. attenuatus Garman 1899, Panama Bay, 1650 m). No species were pre-
viously known from the Galapagos Islands (Grove and Lavenberg 1997).

In this paper, we report the first records of the family from the Galapagos, and describe
Paraliparis darwini and Paraliparis galapagosensis, two new species from upper slope depths. Both
specimens were collected by the Johnson Sea-Link, a three-person research submersible especially
well suited for collecting specimens in topographically complex environments where trawl nets are
unusable.

ISI

S52 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES
Volume 53, No. 11

METHODS

We follow previously published methods of Andriashev and Stein (1998) and Stein et al. (2001)
for counts, measurements, and abbreviations for describing liparids. Proportions are given as % SL
(% HL in parentheses). Terminology of mouth and lower jaw follow Stein et al. (2001). Counts were
obtained using radiographs. Pectoral girdles were removed and then cleared and stained with Alizarin
Red S by the method of Andriashev, Neyelov, and Prirodina (1977).

The following abbreviations for counts and measurements have been used:
For counts

V: vertebrae (abdominal + caudal)

D: dorsal-fin rays

A: anal-fin rays

C: caudal-fin rays

P: pectoral-fin rays (upper lobe rays + notch rays + lower lobe rays)

gr: gill rakers

pe: pyloric caeca

Rad: radials of pectoral girdle
For sensory pores:

io: infraorbital
n: nasal
pm: preoperculo-mandibular
t: temporal
* measurements:
aAf: distance from center of anus to anal-fin origin
bd: maximum body depth
bdA: body depth at anal fin-origin
E: horizontal diameter of eye
go: length of gill opening
HD: maximum head depth
HL: head length
HW: head width
io: interorbital width (between upper margins of eyes)
lj: lower jaw length
LPL: greatest length of lower lobe of pectoral fin
ma: length from mandibular symphysis to center of anus
NL: length of shortest notch ray of pectoral fin
po: postocular head length
preA: preanal-fin length
preD: predorsal-fin length
sn: snout length; tip of snout to anterior margin of eye
SL: standard length
uj: upper jaw length
UPL: greatest length of upper lobe of pectoral fin

Collection abbreviations follow Leviton et al. (1985). Figures are by the junior author. Pectoral
girdle preparations are deposited with the specimens.

STEIN AND CHERNOVA: GALAPAGOS LIPARIDS 153

TAXONOMIC DESCRIPTIONS
Genus Paraliparis Collett, 1878

Paraliparis Collett, 1878:34 (type species Paraliparis bathybii Collett, 1878 by monotypy).
Burke, 1930:154; Andriashev, 1954:464; Cohen, 1968:385; Stein, 1978a:5, 1978b:37; Andriashev,
1986:14; Stein, Chernova, and Andriashev, 2001:360.

DIAGNOSIS. — One pair of nostrils (nostrils single). Ventral sucking disk absent. A single termi-
nal (sensu Andriashev, 1986) or suprabranchial (sensu Burke, 1930) pore present in temporal canal.
Pectoral fin divided into two lobes or not; if present, lower lobe not forming a single filament.
Pseudobranch absent. Coronal pore absent. Barbels or skin flaps on head absent.

Paraliparis darwini n. sp.
Figure |

DIAGNOsIS. — A Paraliparis with teeth in upper jaw absent, in lower jaw uniserial except very
close to symphysis. Teeth in lower jaw blunt canines, oval in cross-section. Chin gelatinous, chin-pore
pair on anterior rather than ventral surface. Head length 21.8% SL, its width 65%, and depth 84%, its
length. Upper jaw 44% HL. Body depth at anal-fin origin 22.1 (102). Lower pectoral-fin lobe 65%
length of upper lobe. P 23 (15+2+6), V 66, C 8. Rad 4 (3+1), round.

HOLOTYPE. — CAS 86576. Female, 131 mm SL, 149 mm TL. “Johnson Sea-Link” Dive 3971,
NE Pacific, Isla Wolf, Galapagos Islands, Ecuador, 637 m, suction collector, 23 November 1995.
Coll. J. E. McCosker et al.

COUNTS AND MEASUREMENTS. — V 66 (12+54), D 58, A 53, P 23 (15+2+6), C 8 (4/4), gr 12, pc
10. HL 21.8, HW 14.1 (65), HD 18.3 (84.2), sn 8.4 (38.6), E3.8 (17.5), po 11.5 (52.6), 10 8.5 (39.3), go
3.8 (17.5), uj 9.5-(43.9), lj 8.5 (39.3), preD 33.5, preA 40, ma 15.6, aAf 22.9, bd 26 (119), bdA 22.1
OZ SUPPL £5:3.:(70)) Nis sor 35% UPL, LPL: 10:7/0r' 70% UPL.

DESCRIPTION. — Head deep at occiput, not compressed, its width 1.3 in its depth. Anterodorsal
contour of head sloping at angle of about 45°, slightly concave above eye. Snout large, deep, rounded,
gelatinous, slightly projecting beyond upper jaw, its length 2.2 times eye diameter. Subrostral fold
deep, well developed, covering upper lip entirely. Nostril with raised rim, not large, horizontal with
lower margin of eye. Eye longitudinally oval, moderately large, not touching dorsal contour of head.
Interorbital 2.2 times eye. Mouth subterminal and small, oral cleft reaching to below anterior margin
of pupil, posterior end of maxilla (under thick soft skin cover) below posterior third quarter of eye.
Lower jaw subterminal. Chin gelatinous, anterior margin prominent beyond lower jaw, chin pores on
its anterior rather than ventral surface. Teeth on upper jaw absent. Teeth on lower jaw primarily
uniserial; in 3 short oblique rows (about 3, 4 and 6 teeth) at symphysis, but posteriorly forming a single
long row of no fewer than 25 teeth on right side of jaw. Teeth small canines, not sharp, not touching
each other; in the uniserial row posterior teeth slightly larger than anterior; in dorsal view (e.g.,
cross-section) teeth oval. Sensory pores less than nostril in diameter; pores on snout with thickened
margins (contoured), in comparatively deep pits of soft tissue and thus look similar to nostril in diame-
ter. Pores 2-6-7-1, postorbital pore present. Chin pores not closely set, interspace about equal to dis-
tance pm,-pm, each ina shallow pit of soft tissue. Gill opening small, equal to horizontal diameter of
eye, entirely above pectoral-fin base. Gill opening located relatively high on side of body, its dorsal
end horizontal with upper margin of eye. Opercular flap small, triangular, its rounded tip closer to
lower end of gill opening and level with upper margin of eye.

Pectoral fin notched, notch rays 2. Uppermost pectoral ray horizontal with upper margin of eye.
Upper lobe not reaching anal-fin origin. Dorsalmost (6th) lower lobe ray longest, lowermost pectoral
ray inserted below anterior third of postocular space. Coracoid not projecting from ventral contour of

154 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES
Volume 53, No. 11

Ficure 1. Paraliparis darwinin. sp. Holotype, CAS 86576, from off Isla Wolf, Galapagos Islands, Ecuador, in 637 m. A: Lower jaw teeth, right lateral view. Scale | mm.

B: Pectoral-fin structure.

STEIN AND CHERNOVA: GALAPAGOS LIPARIDS 155

body. Lower pectoral-fin lobe length 65% that of upper lobe. Right and left lower pectoral-fin lobes
not touching. A transverse, thick, pocket-shaped skin fold present on isthmus anterior to lower lobes.
Radials 3+1, round or rounded, unnotched, R3 distinctly smaller than others. Scapula with slender
helve of moderate length, coracoid with very slender, long helve, base notched ventrally.

Body hump-backed, deep (depth at anal-fin origin about equal to head length). Anus below third
quarter of postocular space. Anus and small genital papilla in a shallow pit, surrounded by soft radial
skin folds. Dorsal contour of body widely rounded. Maximum depth at dorsal-fin origin. Two free
interneurals present between neural spines 4, 5, and 6. First dorsal ray between neural spines 6 and 7.
Dorsal and anal fins not deep: at mid-length of caudal part of body, fin depth less than body depth. In
posterior third of caudal part of body, dorsal fin slightly deeper than anal fin. Dorsal and anal fins over-
lap about half of caudal fin. Gelatinous tissue moderately developed; thicker on snout and chin. Skin
thin, semitransparent, covered with thick mucous layer, opaque after fixation. Free neuromasts not
found. Pyloric caeca digitate, tips sharp, longest ca. 5.3% SL. Ovarian eggs unripe, less than 0.5 mm
in diameter.

In alcohol, body color pale, yellowish brown. Margins of dorsal and anal fins, caudal and pectoral
fins blackish. Ventrally, snout and head, inner subrostral fold, lips and gill opening rosy brown.
Mouth and gill cavities dusky gray. Gill arches pale. Peritoneum ink-black. Stomach, pyloric caeca,
and intestine pale.

DISTRIBUTION. — Known only from the holotype, collected off Isla Wolf, Galapagos Islands.
The specimen was collected from a gently sloping, sand bottom using rotenone and a suction collec-
tor. Also collected during the same dive were Chauliodus sp., Stylephorus chordatus, Symphurus
diabolicus, and some invertebrates.

ETYMOLOGY. — Named after Charles Darwin, describer of the biological diversity of the
Galapagos Island fauna and its significance.

COMPARISONS. — The new species is most similar in counts to P. galapagosensis (below) but
differs most significantly in number of gill rakers (12 vs. 8). Its general aspect is quite different, hav-
ing a much less rounded and longer head and longer abdominal cavity. Specifically, it differs propor-
tionally from P. galapagosensis in head shape (anterodorsal contour at 45°, snout projecting vs. head
evenly rounded, snout not projecting), head depth (84.2 vs. 100% HL), snout length (39 vs. 34% HL),
upper jaw length (43.9 vs. 36.8% HL), and body depth at anus (102 vs. 75% HL). It is also similar to P.
merodontus Stein, Melendez and Kong 1991 from Chile, but differs most distinctly in the number of
vertebrae (66 vs. 62-64), radials (3+1 vs. 3+0), and pyloric caeca (10 vs. 6-8). Proportionally it differs
most in longer head (21.8 vs. 15.1—17.2) and snout (8.4 vs. 4.45.7), greater predorsal-fin length (33.5
vs. 19.3—-26.9), preanal-fin length (40 vs. 34.5-37.6), and body depth (26 vs. 17.3—20.2).

Paraliparis galapagosensis n. sp.
Figure 2

DIAGNOsIS. — A Paraliparis with upper jaw dentition limited to 34 isolated teeth on each side
of symphysis; lower jaw teeth uniserial except very close to symphysis, recurved anteriorly, tips over-
lapping to forma sharp cutting edge. Chin not gelatinous, chin pore pair on ventral surface of chin. HL
20.7% SL, its depth equal to its length. Upper jaw 37% HL. Body depth at anal-fin origin 15.4 (75).
Lower pectoral-fin lobe equal to length of upper lobe. P 22 (15+1+6), V 67, C8. Rad 4 (3+1), round.

HOLOTYPE. — CAS 86737. Female, 94 mm SL, 104 mm TL. “Johnson Sea-Link” Dive 3949,
Cabo Rosa, S of Isla Isabela, Galapagos Islands, Ecuador, 710 m, suction collector, 11 November
1995. Coll. J. E. McCosker et al. Specimen compressed, head strongly flexed ventrally.

COUNTS AND MEASUREMENTS. — V 67 (12+55), D 60, A 55, P 22 (15+1+6), C 8 (4/4), gr 8, pc 9.
HL 20.7% SL, HW - (-), HD 20.7 (100), sn 6.7 (33.7), E 4.1 (20.0), po 11.1 (58), 10 6.7 (32.6), go 3.3
(15.8), uj 7.6 (36.8), lj 8.7 (42.1), preD 30.6 (136.1), preA 33.0 (147.2), ma 13.6 (60.4), aAf 16.3

PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES

156

Volume 53, No. 11

‘AINJONYS Ulf-[BIOJIAg : “WU | B]BIC “MATA [eIO}L] ISL “YIIN} Mel IOMOT :W ‘AT[e.QUDA PoxaT A[SuoINs
peoy ‘possasduroo uourtoadg “wi (| / Ul “1openog ‘spur[s] sosedeyey ‘vjaqes] eIs] JO § ‘esoy OqeD JJo WO *L¢L98 SYD ‘adAjojoH “ds “u szsuasospdnjy3s stupdipavg *7Z PANO

ape TEAR

va Ese

STEIN AND CHERNOVA: GALAPAGOS LIPARIDS 157,

(72.6), bd ca. 27 (108), bdA 15.4 (74.7), UPL 13.0 (63.2), NL - (-), LPL 13.0 (63.2), LPL 100% UPL,
pe 6.5 (41.2).

DESCRIPTION. — Head deep at occiput, anterodorsal contour evenly curved from occiput to
above eye; from there, sloping almost vertically to tip of snout; in life, probably quite compressed.
Snout shallow, rounded, not gelatinous, only slightly projecting beyond upper jaw, short, 1.7 eye di-
ameter. Subrostral fold developed. Nostril with raised rim, not large, about on a horizontal with lower
margin of eye. Eye round, not large, far below dorsal contour of head. Interorbital 1.6 eye. Mouth infe-
rior and small, oral cleft reaching to below anterior margin of eye, posterior end of maxilla (under
thick soft skin cover) below posterior 4 of eye. Lower jaw included. Chin not gelatinous, rounded in
lateral view, sloping gradually posteriorly, chin pores on its ventral surface. Teeth on upper jaw pres-
ent, but only 3-4 isolated teeth on each side of the symphysis. Teeth on lower jaw uniserial except
close to symphysis, forming one long row of no fewer than 40 teeth on right side of jaw. Teeth small,
sharp, recurved anteriorly so that tips overlap to forma sharp cutting edge. Teeth graduated in size, be-
coming increasingly large posteriorly; oval in cross-section. Sensory pores smaller than nostril in di-
ameter; pores on snout without thickened rims (not contoured), not in pits of soft tissue; similar to
nostril in diameter. Pores 2-6?-7-1, postorbital pore not found owing to skin damage, probably pres-
ent. Chin pores not closely set (interspace about equal to distance pm,-pm ), level with chin surface,
not in pits. Posterior to postorbital pore, a row of about 14 free neuromasts present along midline of
body. Gill opening short, 0.8 eye diameter, entirely above pectoral-fin base. Gill opening located rela-
tively high on side of body, level of ventral end well above upper margin of eye. Opercular flap small,
triangular, its tip above middle of gill opening, well above horizontal through upper margin of eye.

Pectoral fin notched, one notch ray. Uppermost pectoral ray above horizontal through upper mar-
gin of eye. Upper lobe reaching anal-fin origin. Longest lower lobe ray fifth from bottom; lowermost
pectoral ray inserted below posterior third of postocular space. Lower pectoral-fin lobe length about
equal to that of upper. The two lower pectoral-fin lobes not touching. Radials 3+1, R3 smallest; all
round, unnotched. Scapula with slender short helve, coracoid with long, very slender helve.

Body strongly hump-backed, deepest just posterior to gill opening. Anterior of vertebral column
strongly curved ventrally to parallel dorsal outline of body, probably natural. Anus below gill open-
ing, almost between insertion of lowest pectoral-fin lobe rays. Ventral contour of body almost
straight. One free dorsal interneural present, insertion of first dorsal ray between vertebrae 6/7. Dorsal
and anal fins not deep: at mid-length of caudal part of body, fin depths less than body depth. In poste-
rior third of caudal part of body, dorsal fin not deeper than anal fin. Dorsal and anal fins overlap first
third of caudal fin. Gelatinous tissue absent. Skin thin, semitransparent. Pyloric caeca digitate, tips
sharply pointed, longest ca. 6.5% SL.

In alcohol, body color pale, yellowish brown. Margins of dorsal and anal fins, caudal and pectoral
fins blackish. Ventrally, snout and head, lips and gill opening rosy brown. Mouth and gill cavities
dusky gray. Gill arches pale. Peritoneum ink black. Stomach, pyloric caeca and intestine pale.

DISTRIBUTION. — Known only from the holotype, collected off Isla Isabela, Galapagos Islands.
The specimen was captured using a suction sampler above a sand bottom with no current. Also col-
lected during the dive were Symphurus diabolicus, Bathypterois sp., Dibranchus erinaceus, and
Rajella eisenhardti, and numerous shrimps and holothurians.

ETYMOLOGY. — The new species is named after the Galapagos Islands of Ecuador, the location
of its collection.

COMPARISONS. — In its counts and tooth arrangement, the new species is most similar to P.
darwini (see above for comparison) although its appearance is quite different. It is also similar to P.
merodontus. from Chile, but differs in number of vertebrae (67 vs. 62—64), anal-fin rays (55 vs.
50-52), radials (3+1 vs. 3+0), and pyloric caeca (9 vs. 6-8). Proportionally it differs most notably in
its greater body depth (ca. 27% SL vs. 17.3—20.2% SL), but it also differs significantly in longer HL

158 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES
Volume 53, No. 11

(20.7 vs. 15.1—17.2) greater predorsal-fin length (30.6 vs. 19.3—26.9), and shorter distance from anus
to anal-fin origin (16.3 vs. 19.4—24.4).

DISCUSSION

Given the recent discoveries of many new snailfishes from the Southern Hemisphere (e.g.,
Andriashev and Stein 1998; Stein et al. 2001), the discovery of new liparid species from the
Galapagos Islands extends the very broad bathymetric and geographic distribution of the family. The
islands are isolated both above and below the water level (McCosker and Rosenblatt 1984; Grove and
Lavenberg 1997) and thus the occurrence of species endemic to the islands, rather than occurring on
the mainland, is “normal.” Estimated ichthyofaunal endemism ts 9.4% of the 437 species reported by
Grove and Lavenberg (1997), the vast majority of which occur at less than 100 m depth, but only 23
species are known from greater depths (ibid.); we suggest that this probably reflects the lack of
deep-water sampling around the islands rather than a depauperate fauna. McCosker and Rosenblatt
(1984) explained the occurrence of endemic species by pointing out that most of the endemics are
from families with low vagility of larvae, 1.e. short larval stages without long pelagic phases. This is
also true of deep-water liparids (Stein 1980). They predicted that “the deep, nearshore steep slopes...
are likely to contain interesting surprises ... modern submersibles . . . allow mankind to reach new
depths .. .” These new species of a family previously unknown from the Galapagos fit this prediction
in location and collection method.

The zoogeographic/systematic relationship between the Galapagos species and the liparid fauna
of the South Pacific is unknown, although the general relationship of the islands’ shallow-water fish
fauna is closest to the eastern tropical Pacific mainland, with nearly 60% of species in common
(McCosker and Rosenblatt 1984; Grove and Lavenberg 1997).

With respect to their tooth arrangement and general morphology, the new species seem most
closely related to P. merodontus from Chile, which has an edentate upper jaw and uniserial lower jaw
teeth, but differs in having 3+0 radials rather than 3+1. Tooth arrangement in both new species seems
to place them in the Paraliparis copei group (see Andriashev 1986, Stein and Andriashev 1990, and
Stein et al. 1991), which is nominally based on all members having teeth uniserial in at least one jaw,
very short or porelike gill opening with reduced opercular flap, and 3 (3+0) pectoral radials. However,
in their pectoral structure and somewhat larger gill opening both differ from P. copei and its subspe-
cies (P. copei gibbericeps Andriashev 1982a, P. copei kerguelensis Andriashev 1982b, and P. copei
wilsoni Richards 1966).

There has been no phylogenetic analysis of the relationships of this group (P. copei copei Goode
and Bean 1896, P. copei gibbericeps, P. copei kerguelensis, P. copei wilsoni, P. merodontus, P.
nassarum Stein and Fitch 1984, P. neelovi Andriashev 1982b, P. paucidens Stein 1978b, P. rosaceus
Gilbert 1890) or thorough examination of their comparative morphology, and indeed, their identity as
a group is somewhat questionable. Pectoral girdle structure is unknown for P. nassarum, P.
paucidens, and P. rosaceus, and as described above, P. darwini and P. galapagosensis differ at least in
their pectoral girdle morphology. These differences suggest that either tooth pattern or pectoral girdle
structure is not a good indicator of relationship. Resolution of this question awaits a full analysis.

ACKNOWLEDGMENTS

The senior author was partially supported by the Office of Oceanic and Atmospheric Research,
National Oceanic and Atmospheric Administration. The junior author was supported by the Russian
Science Foundation, Grants No 99-04-49774, 96-15-97881 and FGP “World Ocean” Project 16. We
thank J. McCosker for bringing the specimens to our attention and for providing his detailed field col-
lection notes.

STEIN AND CHERNOVA: GALAPAGOS LIPARIDS 159

LITERATURE CITED

ANDRIASHEV, A. P. 1954. Fishes of the northern seas of the U. S. S. R. Fauna of the USSR 53. Izdatel’stvo Akad.
Nauk SSSR Moscow [In Russian]. Fishes of the northern seas of the U.S.S.R. Translated 1964. Israel Prog.
Sci. Transl. 836:1—617.

. 1982a. New species of the genus Paraliparis (Liparidae) from the western Antarctic. Report 2. Vopr.

Ikhtiol. 22(2):179—186. [In Russian]

. 1982b. A review of fishes of the genus Paraliparis Collett (Liparidae) from the Kerguelen area,

Subantarctic. Zool. Zh. 61(5):716—725

. 1986. Review of the snailfish genus Paraliparis (Scorpaeniformes: Liparididae) of the Southern Ocean.
Theses Zoologicae 7, Koeltz Scientific Books, Koenigstein. 204 pp.

ANDRIASHEV, A.P. AND D.L. STEIN. 1998. Review of the snailfish genus Careproctus (Liparidae,
Scorpaeniformes) in Antarctic and adjacent waters. Contr. Sci., Nat. Hist. Mus. L. A. County 470:1—63.

ANDRIASHEV, A. P., A. V. NEYELOV, AND V. P. PRIRODINA. 1977. On the method of studying the morphology
and systematics of snailfishes (Liparidae). Zool. Zh. 56 (1):141—147. [In Russian, with English summary |

BurKgE, C. V. 1930. Revision of the fishes of the family Liparidae. Bull. U. S. Nat. Mus. 150:1—204.

CHERNOVA, N. V. AND D. L. STEIN. In Press. Ten new species of Psednos (Pisces, Scorpaeniformes: Liparidae)
from the Pacific and North Atlantic Oceans. Copeia.

COHEN, D. M. 1968. The cyclopterid genus Paraliparis, a senior synonym of Gymnolycodes and Eutelichthys,
with the description of a new species from the Gulf of Mexico. Copeia 1968(2):384-388.

COLLETT, R. 1878. Fiske fra Nordhavs-expeditionens sidste togt, sommeren 1878. Forhandl. Vidensk.-selsk.
Christiania 1878 14:1—106.

GARMAN, S. 1892. The Discoboli. Cyclopteridae, Liparopsidae, and Liparididae. Mem. Mus. Comp. Zool. 14 (pt.
2):1-96.

. 1899. The Fishes. /n Reports on an exploration off the west coasts of Mexico, Central and South Amer-
ica, and off the Galapagos Islands . . . by the U. S. Fish Commission Steamer “Albatross” during 1891 ....
No. XXVI. Mem. Mus. Comp. Zool. 24:1-431, Atlas Pls. 1-85 + A—M.

GILBERT, C. H. 1890. A preliminary report on the fishes collected by the steamer A/batross on the Pacific coast of
North America during the year 1889, with descriptions of twelve new genera and ninety-two new species.
Proc. U. S. Nat. Mus. 13:49-126.

GOopE, G. B. AND T. H. BEAN. 1896. Oceanic Ichthyology, a treatise on the deep-sea and pelagic fishes of the
world. Spec. Bull. U. S. Nat. Mus. (1895):xxxv + 553 pp., with atlas.

GROVE, J. S. AND R. J. LAVENBERG. 1997. The fishes of the Galapagos Islands. Stanford University Press, Stan-
ford. 863 pp.

LeEviTON, A. E., R. H. GIBBS, JR., E. HEAL, AND C. E. DAWSON. 1985. Standards in herpetology and ichthyology.
Part I. Standard symbolic codes for institutional resource collections in herpetology and ichthyology.
Copeia 1985(4):802-832.

McCosker, J. E. AND R. H. ROSENBLATT. 1984. The inshore fish fauna of the Galapagos Islands. Ch. 9 in Key
Environments: Galapagos, R. Perry, ed. Pergamon Press, Oxford. 321 pp.

RICHARDS, W. J. 1966. Paraliparis wilsoni, a new liparid fish from the Gulf of Guinea. Proc. Biol. Soc. Wash.
79:171-174.

STEIN, D. L. 1978a. The genus Psednos a junior synonym of Paraliparis, with a redescription of Paraliparis
micrurus (Barnard) (Scorpaeniformes: Liparidae). Matsya 4:5—10.

. 1978b. A review of the deepwater Liparidae (Pisces) from the coast of Oregon and adjacent waters. Occ.

Pap. Calif. Acad. Sci. 127:1—55.

. 1980. Aspects of reproduction of liparid fishes from the continental slope and abyssal plain off Oregon,
with notes on growth. Copeia 1980(4):687—699.

STEIN, D. L. AND A. P. ANDRIASHEV. 1990. Family Liparididae. Pp. 23 1—255 in Fishes of the Southern Ocean,
O. Gon and P. C. Heemstra, eds. J. L. B. Smith Inst. Ichthyol., Grahamstown. 462 pp., 12 pl.

STEIN, D. L. AND J. E. FITCH. 1984. Paraliparis nassarum n. sp. (Pisces, Liparididae) from off southern Califor-
nia with descriptions of its otoliths and others from north-east Pacific liparidids. Bull. So. Calif. Acad. Sci.
83(2):76-83.

STEIN, D. L. AND L. S. TOMPKINS. 1989. New species and new records of rare Antarctic Paraliparis fishes
(Scorpaeniformes: Liparididae). Ichthyol. Bull. J. L. B. Smith Inst. Ichthyol. 53:18.

160 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES
Volume 53, No. 11

STEIN, D. L., N. V. CHERNOVA, AND A. P. ANDRIASHEV. 2001. Snailfishes (Pisces:Liparidae) of Australia, in-

cluding descriptions of 30 new species. Rec. Austr. Mus. 53:341—406.
STEIN, D. L., R. MELENDEZ C., AND I. KONG U. 1991. A review of Chilean snailfishes (Liparididae,

Scorpaeniformes) with descriptions of a new genus and three new species. Copeia 1991(2):358-373.

PROCEEDINGS
OF THE

CALIFORNIA ACADEMY OF SCIENCES

Volume 53, No. 12, pp. 161-457, 660 figs. October 21, 2002

A Guide to the Lacewings (Neuroptera) of Costa Rica

Edited by

Norman D. Penny
Department of Entomology
California Academy of Sciences, San Francisco, California 94118

PUBLISHED BY THE CALIFORNIA ACADEMY OF SCIENCES
SAN FRANCISCO, CALIFORNIA

PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES

Volume 53, No. 12, pp. 161-457, 660 figs. October 21, 2002

A Guide to the Lacewings (Neuroptera) of Costa Rica

Edited by

Norman D. Penny
Department of Entomology
California Academy of Sciences,
San Francisco, California 94118

Contributing Authors:

Kevin M. Hoffman (Mantispidae)
California Department of Food and Agriculture,
5600 Rickenbacker Road
Bell, California 90201

Martin Meinander (Coniopterygidae)
University of Helsinki,
Helsinki, Finland

Victor J. Monserrat (Hemerobiidae)
Universidad Complutense,
Madrid, Spain

Norman D. Penny
(Ascalaphidae, Berothidae, Chrysopidae, Dilaridae, Polystoechotidae, Sisyridae)

Lionel A. Stange (Myrmeleontidae)
Florida Division of Plant Industry,
P.O. Box 1269
Gainesville, Florida 32602

One hundred and eighty-four species of Neuroptera are currently known from Costa Rica.
Keys to families, subfamilies, tribes, genera and species are presented. For each species di-
agnostic characteristics are stated and illustrated. The general geographical distribution,
and known Costa Rican collection localities are presented for each species. Information is
also given for known elevational ranges, adult flight periods and habitat preferences. One
generic and 13 species synonymies are recognized. There are 29 new combinations.

161

SS
SESS
S ey \

SUL TLZE

art 2 -
2 OR ee on

PENNY: LACEWINGS OF COSTA RICA 163

TABLE OF CONTENTS

eee ees Nees 4, VA PPO Gre ir lek kKik ae 4 6 om ds. ore ignd CORRE cmon Ree 161
SDE TRG sah oe Mae one Nl re Sr tis Or «5 Ree 164
See ee ON TOT VICSL Reve Sls 5 oss ona eee wb e Ae a meee sk MSW MR oe 167
ingiicemacw lf axonomic Combinations. .o....2......... 0.0) nde eine woe te hee 167
Fe Te SS PR WERE MONS ak a ibn Hs we Oe IL A SU BR SWE 168
RRS OLOLIT here eM eT ER ONE Soe ce eis ee oe ee eR es Se ed 168
= LE BEQELSIGINGS ooo. & occa gh Ba eres oe ae Ae a eres ane 169
eee DT MAMET tt.) SN yO os sg ld ED eee tae Be eT Cou oh 169
BoD ATRSS 2. copii ures AAR ee be cr eons ea! Se Ly
anceuneancd)istribution of Newropteta... 2... 2. ec bee ieee ae wee ae es 173
eee Te rec aMe CHIME AL te) MeN es so a gies, cw ol a ed eens PRI oy eos ol LTS
ie AR RCORAT I INICIAL CMM EY MAN BAM Re es Be. cicale ALE eR 2) Re ee ed ety of 176
Pee ERED ACIRT OA Oo) LAL OND. chor Eds 5 S49 aoe web Pee, A OL 186
ce TERS OPEN ToMry a Ne, PRN SS cede, 2s nds. Xie ai, De Ae Se ee 187
MEO MLO CI Ids Fae he da isle, = eve is SR Oe as PO, NO AD. EL 227
ee eee eel tl anitcaomm eye y ee Te at Is 8 Oi cc is a 4s ARR Re ae Ea) eee ad ee oa Wahab 236
STOP OT» SDS RCL OB es Eee sats A ot ie ee one een bee Me kerr ieee eae 238
2 OLDE) i S0CUSy SCENE ee sak oa 2. Aa coe Pe or roe Ae se ae 254
Mame Vviaticle ONtIGAC mir es Me ee ed aa ec ene eL Te wae wee A PATS)
PPM SIOCEMOMGE SR CFE a cies aa cts ad ensue Lome wegen De ee PS SAP 290
PARARNORSGN BURA H A POR ONE nee ce N idee as Gali a fossa dts oA, 40a iw wm whl in oh aha 290
Ergo A AMAGIIS 7. 2 oY MR TN MND ce eho ea wlis Sa late a ws a wn oe OR NE aR 292
eee Cen ee en a Pet rs). oz cs ow a wpa eetar ees See POSE eT 293
2S 22 VST ETDS TUS cy ieee seeing oid apt ich ieael 450
an ied ne ROO LUNE PO eM en Pee ote oot wi dx Space 4 ay'sys Wiis 'oce. oa ware Eee Loa ttod 451

DEPT TAS Mt ca lt Net es nes 8s ety ese 454

PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES

Volume 53, No. 12

LIST OF INCLUDED TAXA

ASCALAPHIDAE
Haplogleniinae

1. Amoea vacuus (Gerstaecker, 1893)

2. Ascalobyas albistigma (Walker, 1853)

3. Ascalobyas microcerus Rambur, 1842

4. Haploglenius luteus Walker, 1853

5. Haploglenius peruvianus Weele, 1908

6. Neohaploglenius flavicornis (McLachlan, 1871)
Ascalaphinae

7. Ameropterus breviantennis Penny, 1982

8. Ameropterus consors (Gerstaecker, 1893)

9. Ameropterus mexicanus (Weele, 1908)

10. Ameropterus subripiens (Walker, 1853)

11. Ameropterus trivialis (Gerstaecker, 1888)

12. Cordulecerus inquinatus Gerstaecker, 1888

13. Cordulecerus praecellens (Gerstaecker, 1884)

14. Cordulecerus subiratus (Walker, 1853)

15. Ululodes bicolor (Banks, 1895)

16. Ululodes cajennensis (Fabricius, 1787)

17. Ululodes macleayanus (Guilding, 1825)

BEROTHIDAE

18. Spiroberotha sanctarosae Adams, 1990

CHRYSOPIDAE
Apochrysinae
19. Domenechus mirifica (Gerstaecker, 1888)
20. Loyola croesus (Gerstaecker, 1893)
Chrysopinae - Leucochrysini
21. Berchmansus elegans (Guérin-Ménéville, 1838)
22. Gonzaga palliceps (McLachlan, 1867)
23. Gonzaga torquatus Navas, 1913
24. Leucochrysa (Leucochrysa) adamsi Penny, 2001
25. Leucochrysa (L.) clara (McLachlan, 1867)
26. Leucochrysa (L.) colombia (Banks, 1910)
27. Leucochrysa (L.) lestagei Navas, 1922
28. Leucochrysa (L.) magnifica (Banks, 1920)
29. Leucochrysa (L.) pretiosa Banks, 1910
30. Leucochrysa (Nodita) amazonica Navas, 1913
31. Leucochrysa (N.) amistadensis Penny, 2001
32. Leucochrysa (N.) antica Navas, 1913
33. Leucochrysa (N.) askanes (Banks, 1946)
34. Leucochrysa (N.) caucella (Banks, 1910)
35. Leucochrysa (N.) compar (Navas, 1921)
36. Leucochrysa (N.) cornesta Banks, 1944
37. Leucochrysa (N.) cortesi Navas, 1913
38. Leucochrysa (N.) duarte Banks, 1946

39. Leucochrysa (N.) egregia Navas, 1913

40. Leucochrysa (N.) forcipata Penny, 1998

41. Leucochrysa (N.) indiga (Navas, 1928)

42. Leucochrysa (N.) laertes (Banks, 1946)

43. Leucochrysa (N.) lancala Banks, 1944

44. Leucochrysa (N.) lenora (Banks, 1944)

45. Leucochrysa (N.) navasi Kimmins, 1940

46. Leucochrysa (N.) nigrovaria (Walker, 1853)

47. Leucochrysa (N.) oenops Adams, 1987

48. Leucochrysa (N.) pallescens Banks, 1946

49. Leucochrysa (N.) postica Navas, 1913

50. Leucochrysa (N.) ratcliffei Penny, 2000

51. Leucochrysa (N.) serrei (Navas, 1923)

52. Leucochrysa (N.) superior Navas, 1913

53. Leucochrysa (N.) virginiae Penny, 1998

54. Vieira leschnaulti (Navas, 1911)
Chrysopinae - Belonopterygini

55. Nacarina cordillera (Banks, 1910)

56. Nacarina megaptera (Navas, 1927)

57. Nacarina titan Banks, 1915
Chrysopinae - Chrysopini

58. Ceraeochrysa acmon Penny, 1998

59. Ceraeochrysa arioles (Banks, 1944)

60. Ceraeochrysa berlandi (Navas, 1924)

61. Ceraeochrysa caligata (Banks, 1946)

62. Ceraeochrysa cincta (Schneider, 1851)

63. Ceraeochrysa claveri (Navas, 1911)

64. Ceraeochrysa costaricensis Penny, 1997

65. Ceraeochrysa cubana (Hagen, 1861)

66. Ceraeochrysa defreitasi Penny, n.sp.

67. Ceraeochrysa discolor (Navas, 1914)

68. Ceraeochrysa elegans Penny, 1998

69. Ceraeochrysa everes (Banks, 1920)

70. Ceraeochrysa gradata (Navas, 1913)

71. Ceraeochrysa inbio Penny, 1997

72. Ceraeochrysa nigripedis Penny, 1997

73. Ceraeochrysa pseudovaricosa Penny, 1998

74. Ceraeochrysa sanchezi (Navas, 1924)

75. Ceraeochrysa smithi (Navas, 1914)

76. Ceraeochrysa tauberae Penny, 1997

77. Ceraeochrysa valida (Banks, 1895)

78. Chrysoperla exotera (Navas, 1913)

79. Chrysoperla externa (Hagen, 1861)

80. Chrysopodes (Chrysopodes) costalis (Schneider,

1851)

81. Chrysopodes (C.) nevermanni Navas, 1928

82. Chrysopodes (C.) varicosa Navas, 1914

PENNY: LACEWINGS OF COSTA RICA

83. Chrysopodes (C.) victoriae Penny, 1998
84. Chrysopodes (Neosuarius) collaris (Schneider,
1851)

85. Chrysopodes (N.) crassinervis Penny, 1998
86. Chrysopodes (N.) crassipennis Penny, 2000
87. Chrysopodes (N.) oswaldi Penny, n.sp.

88. Meleoma poolei Adams, 1969

89. Meleoma titschaki Navas, 1928

90. Plesiochrysa brasiliensis (Schneider, 1851)
91. Plesiochrysa elongata (Navas, 1913)

92. Ungla pallescens Penny, 1998

CONIOPTERYGIDAE
Aleuropteryginae

93. Aleuropteryx clavicornis Meinander, 1995

94. Aleuropteryx rugosa Meinander, 1995

95. Neoconis dentata Meinander, 1972

96. Neoconis marginata Meinander, 1972
Coniopteryginae

97. Coniopteryx (Xeroconiopteryx) falcata Meinander,

1995

98. Coniopteryx (Scotoconiopteryx) cyphodera John-
son, 1978

99. Coniopteryx (S.) flinti Meinander, 1975

100. Coniopteryx (S.) fumata Enderlein, 1907

101. Coniopteryx (S.) fumicolor Meinander, 1972
102. Coniopteryx (S.) isthmicola Meinander, 1972
103. Coniopteryx (S.) panamensis Meinander, 1974
104. Coniopteryx (Coniopteryx) californica
Meinander, 1974

105. Coniopteryx (C.) freytagorum Johnson, 1978
106. Coniopteryx (C.) mexicana Meinander, 1974
107. Semidalis arnaudi Meinander, 1972

108. Semidalis manausensis Meinander, 1980

109. Semidalis mexicana Meinander, 1972

110. Semidalis panamensis Meinander, 1974

111. Semidalis rondoniensis Meinander, 1982

112. Semidalis soleri Monserrat, 1985

DILARIDAE

113. Nallachius americanus (McLachlan, 1880)
114. Nallachius parkeri Penny, 1994

115. Nallachius pulchellus Banks, 1938

HEMEROBIIDAE

116. Hemerobius bolivari Banks, 1910
117. Hemerobius discretus Navas, 1917
118. Hemerobius edui Monserrat, 1991
119. Hemerobius gaitoi Monserrat, 1996

120. Hemerobius hernandezi Monserrat, 1996
121. Hemerobius hirsuticornis Monserrat, 1999
122. Hemerobius jucundus Navas, 1927

123. Hemerobius martinezae Monserrat, 1996
124. Hemerobius nigridorsus Monserrat, 1996
125. Hemerobius pennii Monserrat, 1996

126. Hemerobius withycombei (Kimmins, 1928)
127. Megalomus minor Banks, 1905

128. Megalomus pictus Hagen, 1861

129. Micromus subanticus (Walker, 1853)

130. Notiobiella mexicana Banks, 1913

131. Notiobiella rubrostigma Navas, 1914

132. Nusalala brachytera Oswald, 1996

133. Nusalala championi Kimmins, 1937

134. Nusalala colombiensis (Banks, 1910)

135. Nusalala ghioi Monserrat, 2000

136. Nusalala irrebita (Navas, 1929)

137. Nusalala tessellata (Gerstaecker, 1888)
138. Nusalala uncata Kimmins, 1937

139. Nusalala unguicaudata Monserrat, 2000
140. Sympherobius angustus (Banks, 1904)

14
142. Sympherobius similis Carpenter, 1940

. Sympherobius arizonicus Banks, 1911

143. Sympherobius subcostalis Monserrat, 1998

MANTISPIDAE
Calomantispinae
144. Nolima infensus Navas, 1924
Mantispinae
145. Buyda phthisica (Gerstaecker, 1885)
146. Climaciella brunnea (Say, 1824)
147. Climaciella obtusa Hoffman, n.sp.
148. Climaciella porosa Hoffman, n.sp.
149. Dicromantispa debilis (Gerstaecker, 1888)
150. Dicromantispa gracilis (Erichson, 1839)
151. Dicromantispa sayi (Banks, 1897)
152. Dicromantispa synapsis Hoffman, n.sp.
153. Entanoneura batesella (Westwood, 1867)
154. Haematomantispa nubeculosa (Navas, 1933)
155. Leptomantispa chaos Hoffman, n.sp.
156. Leptomantispa nymphe Hoffman, n.sp.
157. Leptomantispa pulchella (Banks, 1912)
158. Zeugomantispa compellens (Walker, 1860)
159. Zeugomantispa minuta (Fabricius, 1775)
160. Zeugomantispa viridula (Erichson, 1839)
Symphrasinae
161. Plega hagenella (Westwood, 1867)
162. Trichoscelia remipes (Gerstaecker, 1888)

165

166 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES
Volume 53, No. 12

MYRMELEONTIDAE 173. Elachyleon punctipennis Esben-Petersen, 1927
Myrmeleontinae - Acanthaclisini 174. Eremoleon dunklei Stange, 1999

163. Vella fallax (Rambur, 1842) 175. Eremoleon genini (Navas, 1924)
Myrmeleontinae — Brachynemurini 176. Glenurus heteropteryx Gerstaecker, 1885

164. Dejuna fenestrata (Banks, 1913) 177. Glenurus luniger Gerstaecker, 1893

165. Dejuna mimica (Stange, 1970) 178. Glenurus proi Navas, 1929

166. Dejuna setosa (Stange, 1970) 179. Purenleon abruptus Stange, n.sp.
Myrmeleontinae - Myrmeleontini 180. Purenleon debilis (Gerstaecker, 1893)

167. Myrmeleon (Nehornius) perspicuus Gerstaecker, 181. Purenleon iniquus (Navas, 1914)

1893 182. Purenleon parallelus (Banks, 1935)

168. Myrmeleon timidus Gerstaecker, 1888

169. Myrmeleon uniformis Navas, 1920 POLYSTOECHOTIDAE
Myrmeleontinae - Nemoleontini 183. Polystoechotes punctatus (Fabricius, 1787)

170. Dimarella angusta (Banks, 1908)

171. Dimarella garciai (Navas, 1932) SISYRIDAE

172. Dimarella mixteca Miller, 1989 184. Sisyra apicalis Banks, 1908

PENNY: LACEWINGS OF COSTA RICA

INCLUDED NEW SYNONYMIES
Note: New synonymies are indicated in the body of the text with bold face type.

GENUS
Diazus Navas, 1914 = Purenleon Stange, new name

SPECIES

Haploglenius angulatus Gerstaecker, 1894 = Neohaploglenius flavicornis (McLachlan, 1871)
Leucochrysa luctuosa Banks, 1914 = Leucochrysa (Nodita) egregia Navas, 1913

Chrysopa ceratica Navas, 1911 = Leucochrysa (Nodita) nigrovaria (Walker, 1853)

Nodita maculata Navas, 1928 = Leucochrysa (Nodita) nigrovaria (Walker, 1853)

Chrysopa indicata Navas, 1914 = Ceraeochrysa gradata (Navas, 1913)

Chrysopa infausta Banks, 1946 = Ceraeochrysa gradata (Navas, 1913)

Mantispa limbata Gerstaecker, 1885 = Entanoneura batesella (Westwood, 1867)

Mantispilla flavicornis Navas, 1930 = Zeugomantispa minuta (Fabricius, 1775)

Mantispilla rubricata Navas, 1924 = Zeugomantispa minuta (Fabricius, 1775)

Mantispilla viridata Navas, 1924 = Zeugomantispa minuta (Fabricius, 1775)

Mantispa viridis Walker, 1853 = Zeugomantispa minuta (Fabricius, 1775)

Myrmeleon leptaleus Navas, 1924 = Myrmeleon uniformis Navas, 1920

Elachyleon punctipennis pulchellus Esben-Petersen, 1933 = El. punctipennis Esben-Petersen,
1927

INCLUDED NEW TAXONOMIC COMBININATIONS
Note: New taxonomic combinations are indicated in bold face type in the text.

Chrysopidae

Gonzaga palliceps (McLachlan, 1867)
Leucochrysa (Nodita) duarte (Banks, 1946)
Mantispidae

Buyda phthisica (Gerstaecker, 1885)
Dicromantispa debilis (Gerstaecker, 1888)
Dicromantispa gracilis (Erichson, 1839)
Dicromantispa sayi (Banks, 1897)
Entanoneura batesella (Westwood, 1867)
Haematomantispa nubeculosa (Navas, 1933)
Leptomantispa pulchella (Banks, 1912)
Zeugomantispa compellens (Walker, 1860)
Zeugomantispa minuta (Fabricius, 1775)
Zeugomantispa viridula (Erichson, 1839)
Myrmeleontidae

Dejuna fenestrata (Banks, 1913)

Dejuna mimica (Stange, 1970)

Dejuna setosa (Stange, 1970)

Purenleon albovaria (Banks, 1942)
Purenleon bistictus (Hagen, 1861)
Purenleon clavatus (Navas, 1914)
Purenleon connexus (Banks, 1920)

168 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES
Volume 53, No. 12

Purenleon cubensis (Alayo, 1968)
Purenleon debilis (Gerstaecker, 1894)
Purenleon imbellus (Banks, 1941)
Purenleon iniquus (Navas, 1914)
Purenleon inscriptus (Hagen, 1861)
Purenleon minor Banks, 1927
Purenleon nubipennis (Navas, 1917)
Purenleon parallelus Banks, 1935
Purenleon reductus Banks, 1941
Purenleon zayasi Alayo, 1968

PREFACE

Biologists and administrators in Costa Rica are in the midst of an exciting project to inventory the
biological diversity of their country. This is not an easily achievable mission. It will require the com-
bined efforts of hundreds, perhaps thousands, of taxonomists from many disparate disciplines willing
to devote their time and energy to supporting this task. At this time we do not know if all groups of
plants and animals will ever be completely documented, and if so, how long it will take. However, in
order to accomplish a complete inventory within a reasonable time frame, each group of specialists
must proceed with the idea in mind that they are providing a small portion of that information. The
California Academy of Sciences, through its cadre of biologists and scientific publication system, is
proud to support this ambitious goal. The following publication is the result of one of these efforts. It is
hoped that this work will provide the stimulus for more monographs on the fauna and flora of Costa
Rica, as well as lay the foundation for further studies of Neuroptera in the region.

Until recently the term Neuroptera was applied collectively to a group of insects that are now gen-
erally recognized as three orders: Raphidioptera (snakeflies), Megaloptera (dobsonflies and
fishflies), and Neuroptera in a more restricted sense (lacewings, antlions, owlflies, dustywings, and
their allies). This monograph treats Neuroptera only in the more restricted definition.

INTRODUCTION

From the first Spanish explorers to most recent biologists, Costa Rica has continued to impress
people as a “costa rica,” or rich coast. For biologists and tourists alike, the country’s plants and ani-
mals are amazing for their diversity as well as their colorful and bizarre forms. From the iridescent Re-
splendent Quetzal (Pharomachrus mocinno) and Turquoise Cotingas (Cotinga ridgwayi) to brilliant
Poison-Arrow Frogs (Dendrobates spp.) and Golden Toads (Bufo periglenes), from metallic blue
Morpho spp. butterflies and golden yellow Chrysina resplendens scarab beetles to large pink Cattleya
skinneri orchids, bright coral reef fishes and brilliantly colored sea slugs, it is amazing that a country
so small (only 51,054 sq. km, see Map 1, 2) can contain so many biological wonders. Costa Rica, the
size of West Virginia, has 820 bird species (more than all of North America). There are 1100 species
of orchids (Walter 1983). The number of butterfly species (543) is much greater than that ofall of Brit-
ain and Europe (338) (Higgins and Riley 1970). The number of species of butterflies is also equal or
greater than that of some well-studied tropical countries, such as Malaysia, Liberia, and Panama
(DeVries 1986). This biodiversity extends also to the Neuroptera. When the present study was initi-
ated, only 40 species were known in the literature from Costa Rica. Now, we know of 184 species and
there are almost certainly additional species that we have not yet seen. This compares favorably with
the 243 species known from all of Europe, or the 200+ species known from the lowland Amazon Ba-
sin.

PENNY: LACEWINGS OF COSTA RICA 169

BIOGEOGRAPHY

The Neuroptera fauna of Costa Rica display several distinctive distributional patterns that reflect
climatic/floristic zones. Guanacaste Distribution. Guanacaste Province is a relatively dry climatic
zone where a number of species reach their southern limits, or are endemic to the area. These include
such species as Notiobiella mexicana Banks and Spiroberotha sanctarosae Adams.

A second pattern is a Pacific Coastal Distribution, with species found all along the Pacific low-
lands and central valley, but not on the Caribbean slopes or lowlands. Species displaying this distribu-
tion include Micromus subanticus and Vella fallax.

A third group of species is found in the Caribbean lowlands, usually also distributed into north-
ern Heredia Province. Species that show this pattern are frequently present further south in Panama
and South America, but not further north. Some examples are Buyda phthisica and Leucochrysa
(Leucochrysa) pretiosa.

A fourth pattern is a mid- to high-elevational Distribution along the high central mountains.
These species are also typically found in South America and reach their northern limit in Central
America. Hemerobius bolivari 1s an example of this type of distribution, being found from central
Chile to Guatemala. The reverse of this pattern is found in Polystoechotes punctatus. It ranges from
Canada and northern U.S.A., south through the mountainous areas of western U.S.A. and Mexico to
Costa Rica and northern Panama.

A further element may be the endemic fauna. These include the already mentioned lowland spe-
cies in Guanacaste Province, and several upland species in various mountain ranges. There are many
species included in this monograph which at this time are known only from Costa Rica. However, be-
cause this country is much better studied than some neighboring countries, it may be premature to call
any Neuroptera species endemic to Costa Rica until further collecting is done.

In Guanacaste Province several species of Hemerobiidae known from further north can be found
at low elevations, such as Micromus subanticus and Megalomus minor. However, in the central and
southern parts of Costa Rica, hemerobiids have not yet been found at low elevations. At
mid-elevations (1300-1600 m), such as Monteverde and Las Alturas, the dominant species appears to
be the South American Hemerobius bolivari.

Many species are found extensively throughout North and Central America, and the land forms of
Costa Rica appear not to have formed permanent barriers to movement of these species. Such species
include the strong-flying owlfly Ululodes macleayanus, as well as smaller ubiquitous species com-
mon to agricultural habitats, like Chrvsoperla externa, or forest edge habitats, like Megalomus minor,
Mantispa viridula and Climaciella brunnea.

Finally, a note of caution should be exercised when looking at geographical distributions of
Neuroptera, both within Costa Rica and regionally. Many of these insects are small and not often col-
lected using commonly utilized trapping techniques. Their currently recognized distributions are not
yet well known and many may be far more extensive than presently known. Some species known from
a few widely separated collection sites may eventually be found throughout Central America. We are
fortunate in having so many recent records from Costa Rica. The data presented herein will give some
guidance as to distribution, but habitat and elevational data may also provide help in finding these spe-
cies in other parts of the country, and in neighboring countries, where they have not yet been collected.

SCOPE OF THIS MANUAL

Inclusions: This manual is not meant to be a definitive monograph of the Neuroptera fauna of
Costa Rica. Rather, it is meant to provide guidance in identifying species within this fauna, as an aid to
further studies of biology, ecology, and pest management. Although coniopterygids and some
chrysopids and hemerobiids cannot be identified without using male genitalia, most neuropterans in

Volume 53, No. 12

PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES

170

BIQUIO|OD ;

ry
/ ud

eas ueeqqueo

“pOLIOUry [e.yuaD WaYyINOs Jo dey “| dv

\
\*\\)

enbeieoin

PENNY: LACEWINGS OF COSTA RICA 171

\\i Lake

aes AY monnacun

j CARIBBEAN

eed

: Heredia *.

Alajuela

Guanacaste

wridiceaas
San Jose. Cartago
Limon
_ San™
‘... Jose

Puntarenas
PACIFIC OCEAN

Map 2. Map of Costa Rica.

172 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES
Volume 53, No. 12

Costa Rica can be identified using non-genitalic characters such as wing venation and head markings.
However, this manual must be used with caution, as there are almost certainly some species in Costa
Rica which have not yet been collected or described. If there is any doubt about the identity of a spe-
cies, consultation of appropriate specialists or more detailed literature is recommended.

Exclusions: The authors have only treated the fauna based on adult material, except in the
Myrmeleontidae. Kady and Maurice Tauber (Cornell University, Ithaca, New York, U.S.A.) have
reared many larval chrysopids from Costa Rica and Sergio de Freitas (Universidade Estadual de Sao
Paulo, Jaboticabal, Brazil) has done similar work in Sao Paulo State, Brazil. We encourage them and
others to continue to provide insights into the larval stages of the Costa Rican neuropteran fauna.

Non-Costa Rican Families. The authors have examined thousands of specimens for the current
work and have documented the diversity of Neuroptera within Costa Rica as completely as is possible
at the present time. Three other New World families may eventually be found within the country.
They are:

Ithonidae. Until recently this family was only known from Australia, the high mountains of
southeast Asia, Mexico and the deserts of southwestern U.S.A. However, Penny (1996) described a
new genus and species of ithonid from the high mountains of northern Honduras. Males of the Hondu-
ras species are weak fliers and females are flightless, making dispersal difficult. Because high altitude
populations generally are localized and inaccessible, they are seldom adequately sampled and it is
possible that members of Ithonidae may subsequently be found in the higher mountainous regions of
Costa Rica.

Nemopteridae. In the New World this family was known for many years from two species of
Crocinae in Argentina and one species of Nemopterinae in Chile. Knowledge of their geographical
distribution was expanded when Stange and Williner (1981) found a species of Crocinae in Bolivia,
and again when Mansell (1983) encountered a new genus of Crocinae in southern Peru. Subsequently,
Miller and Stange (1989b) have collected the genus described by Mansell along the Caribbean coast
of Venezuela. Finding Nemopteridae so far north was very unexpected, and has fueled much specula-
tion as to the actual distribution of nemopterids in the Neotropics. Crocinae are relatively small and
transparent-winged, and most are cave-dwellers. Almost all specimens are picked up by a few special-
ists who recognize this subfamily and know where to look for them, most often rearing them from lar-
vae.

Osmylidae. In the Western Hemisphere osmylids are known only from South America. This
family is frequently collected in the Andes of Chile and species of /sostenosmylus and Gumilla are
found in the mountains of Santa Catarina State in southern Brazil. North of Chile they have been less
frequently collected in the Andes as far north as Ecuador. Larvae most often live in mosses along the
margins of mountain streams. There appears to be no barrier to their dispersal along the Andes into
Colombia, although they have never been recorded from that country. The question remains as to
whether the backbone of mountains into Central America is high enough and has been continuous
enough over geological time to allow survival of osmylids in Costa Rica.

ABUNDANCE

Perhaps no feature of the Costa Rican Neuroptera fauna is so striking as the relatively large num-
ber of species which are seldom collected. Despite having wide geographical distributions and ap-
pearing to be present in the adult stage during most, if not all of the year, these species have eluded the
best efforts of researchers and parataxonomists alike to capture them. For example, the very large and
unusual dark-pigmented green lacewing, Loyola croesus, which is found from southern Mexico to
Peru has only been collected in Costa Rica three times. Relatively few species can be collected with
regularity and predictability. Only by gaining a much better knowledge of individual species’ life his-
tories can we begin to understand the role that these predators play in the Costa Rican ecosystem. It is

PENNY: LACEWINGS OF COSTA RICA 173

hoped that the present monograph will create a foundation that students can use to undertake the stud-
ies needed to better understand this role.

COLLECTING AND DISTRIBUTION OF NEUROPTERA

Two methods have proven most effective for collecting Neuroptera—beating vegetation with a
sweep net and light trapping. Two additional methods that have also been moderately effec-
tive—flight intercept traps (such as Malaise traps) and molasses bait traps (for Chrysopidae).

Three families of Neuroptera are taken regularly by beating vegetation: Coniopterygidae,
Hemerobiidae, and Chrysopidae. Coniopterygidae are usually found in dense forest vegetation low to
the ground. They are generally found in the 0-1 m height range. Occasionally, however, they are
found in more open, sunny habitats.

Coniopterygidae are most active, and most easily seen, very early in the morning (about 6:00 to
8:00 a.m.). At that time they frequently can be seen fluttering across forest trails, or hovering near
dense vegetation. In the early morning the white powder of their wings shows clearly in flight against
the dark forest backdrop.

Hemerobiidae can be taken by beating tree branches at about 2 to 3 m height in primary forests.
They are difficult to see because of their small size and pale brown coloration. When fluttering to
neighboring branches, they resemble small moths that frequent the same habitat. They often prefer
resting on dark brown surfaces, such as dead leaves, vines, or tree trunks, although they are also often
found under green leaves. Often when a branch is beaten, brown lacewings will not fly at all, but rather
drop to the forest floor and lie quiescent for several seconds. As tropical forest leaves often bear a
heavy load of debris, rapping ona branch will send a shower of debris cascading to the forest floor, and
hemerobiids generally go unseen with this debris when feigning death. When specifically looking for
Hemerobiidae, it is best to use a beating sheet, or to hold a collecting net open below the branch and
then carefully look through the debris for individual brown lacewing specimens.

Hemerobiidae have also been taken effectively with light traps at low elevations in Guanacaste
Province and at mid-elevations in Escazi and San Vito. The highest elevation recorded for
Hemerobiidae in Costa Rica is a specimen of Hemerobius bolivari from 2360 m elevation in Zona
Protectora Las Tablas, near Cerro Echandi.

Chrysopidae appear to have a somewhat narrower elevational range than Hemerobiidae, being
found from sea level to ca. 1900 m. Leucochrysine green lacewings are usually collected in the same
way as brown lacewings, by beating understory branches at 2-3 m height. Leucochrysini have less of
a tendency to feign death, and more frequently are seen fluttering to nearby branches. They seem to
prefer woody dicot understory saplings. Best results have been obtained by beating young trees near
streams in primary forest, where humidity is high and light intensity low. The only Chrysopini found
in primary forest have been Chrysopodes species. Although no habitat information is available for
Costa Rican Apochrysini, in other parts of the Neotropics (Brazil) and Papua New Guinea this tribe
has only been taken in primary forest. Chrysopini can often be taken in large numbers in open pastures
or grasslands. The dominant species appears to be Chrysoperla externa, which, as adults, feed on
grass pollen. This species can be found even in isolated pastures deep in forest with few or no grasses,
indicating excellent powers of dispersal.

Fewer species of Chrysopidae and Hemerobiidae seem to be found in young secondary forest, al-
though Chrysopodes seems to do well in such habitats.

An effective method for collecting most groups of Chrysopidae is light trapping. Both mercury
vapor and ultra-violet lights, as well as ordinary fluorescent and incandescent bulbs attract some spec-
imens of Chrysopidae. One of only three known specimens of Costa Rican Loyola croesus was at-
tracted to lights (Luis Diego Gomez, personal communication). However, chrysopids have at times
been seen flying past bright lights at night, with little apparent attraction. When attracted to lights, they

174 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES
Volume 53, No. 12

often prefer to rest on the undersurfaces of overhanging nearby objects, such as eaves of buildings and
leaves of nearby trees.

Other groups of Neuroptera, such as Ascalaphidae, Myrmeleontidae, Berothidae, and
Mantispidae are most frequently collected with light traps.

Berothidae are apparently restricted to Guanacaste Province and emerge during the driest part of
the year, from December to March.

Mantispidae are common in forested and forest-edge areas of Costa Rica, and can occasionally be
collected with a net. However, they seem to be collected most frequently at lights. The exception to
this is the wasp-mimicing species, Climaciella brunnea, which is seen most often in open meadows
and pastures on clumps of flowers, such as umbels and composites. They imitate wasps very closely
(Opler 1981), and make no attempt to flee quickly or hide. In the United States, this species often has
short-duration mass emergences, where many individuals will be seen on numerous flowers in an
area, only to disappear completely a week later (Jean Laffoon, personal communication). Mantispidae
are most abundant in lowland areas of Costa Rica and have not been collected above 1500 m. Buyda
phthisica and a closely-related new species from Panama have been taken only in lowlands on the Ca-
ribbean side of the central mountain ranges.

Sisyridae should be found in many parts of Costa Rica along the margins of streams where fresh-
water sponges are found, but have to date only been collected once within the country, at a light trap
near Canas in Guanacaste Province.

Ascalaphidae are active at dusk, skimming over the soil surface in open areas and among the tree
top canopies in forests. They are difficult to see in failing light, and even harder to catch. They are very
strong fliers, quickly pursuing small flying insects at twilight. They sometimes hit sheets and other
barriers at light traps in their pursuit of insects attracted to the lights. Some species of owlflies in the
southwestern U.S.A. (for example, Ululodes arizonensis Banks) do not seem to respond to the light it-
self, and if not captured quickly, will hit the light sheet while pursuing insects, reorient themselves in
moments, and quickly fly away. Other species in northern Brazil (for example, Ameropterus
delicatulus (McLachlan)) appear to be strongly attracted to lights and will stay at a light for hours.

The Haplogleniinae appear to be forest dwellers, while most Ascalaphinae are found in more
open situations. Larval Haplogleniinae occasionally can be seen waiting motionlessly on understory
vegetation. They are extremely flattened and appear very similar to the fungal growths that often spot
tropical leaves. It is suspected that these larvae have multi-year life cycles. Haplogleniine adults occa-
sionally can be seen resting singlely on low vegetation in the forest with their wings open. Larvae of
Ascalaphinae rest on the soil surface and blend in so well with their substrate that they are seldom
seen. Adult Ascalaphinae rest during the day on twigs with their wings folded along the branch so that
they are only seen infrequently. The ascalaphine genus Cordulecerus is an exception, being collected
most often in the forest, where they are sometimes found aggregating in large clusters, especially near
streams (Hogue and Penny 1989).

Myrmeleontidae is the most speciose family of Neuroptera in the world, but its species prefer
drier areas of the world, and are not very diverse in Costa Rica. The greatest diversity of
Myrmeleontidae in the country appears to be in the drier Guanacaste Province. In Costa Rica, only lar-
vae of the genus Myrmeleon make funnels, which are ubiquitous under buildings and bridges
throughout the country below 1500 m elevation. Other genera have larvae which live on the soil sur-
face, just under the soil surface, on ceilings of caves, in tree holes, or in rock crevices. Forest-dwelling
species, such as Eremoleon spp., are not as well known as species that frequent open dry areas, and we
may eventually find a rich, but rarely encountered, forest fauna of ant-lions. Although many individu-
als of forest-dwelling species may be present in any lowland area of the country, they are seldom seen
except at light traps. Although the funnels of Myrmeleon larvae are very common, the adults are more
rarely encountered.

PENNY: LACEWINGS OF COSTA RICA 175

Dilaridae is another family whose members are almost never seen without using light traps. In the
United States, larvae have been found in rotting tree trunks, feeding on wood-dwelling insects. A
study in a Brazilian Amazon forest (Penny and Arias 1981) found adults far more actively attracted to
lights in the forest canopy than at ground level, and adults appeared to be most abundant during the
driest period of the year.

Individuals of Polystoechotidae are seldom encountered in Costa Rica. They have been collected
at mid-elevations at Monteverde and Cartago. In the United States Polystoechotes punctatus is known
to fly at very low temperatures (5° C) and is strongly attracted to smoke. Many have been collected
around camp fires. Polystoechotids are also taken at lights in remote areas. Their geographical range
appears to be shrinking dramatically, perhaps in response to lights in developed areas.

SYSTEMATIC TREATMENT

KEY TO FAMILIES OF COSTA RICAN NEUROPTERA
(after Henry et al. 1991)

1. Forewing length 4 mm or greater; body and wings not covered with a whitish powder ................ 2
1’. Forewing length 3 mm or less; body and wings covered with a whitish powder (Fig. 328) ..... . Coniopterygidae

2(1). Antennae clubbed or knobbed; abdomen long and slender, somewhat resembling Odonata in appearance (Fig. 1);

[SaeRW LOMO DUSEMCUIVCUNAWS! ait cee ein os ters ee ees cao Gn ae emer nice. Gok Beet ne eon ee ete eee Te 3
2'. Antennae filiform, moniliform, or pectinate, not clubbed or knobbed; not resembling Odonata (Fig. 36); larvae with
fitinastralchtonsliohthvicinvedyjawsts «dle eteee 4 BU Sleet ORE A Se ee 4

3(2). An elongate cell present behind point of fusion of Sc and R, in distal part of wing (Figs. 617-622); antennae apically

swollen, but not knobbed, and not longer than the head and thorax combined (Fig. 593) ....... Myrmeleontidae
3’. No elongate cell present behind point of fusion of Sc and R, (Figs. 6-11); antennae apically knobbed, usually as long
as entire body, but sometimes as short as head and thorax combined (Fig. 1). .............. Ascalaphidae

4(2'). Prothorax elongate; forelegs greatly enlarged, raptorial (Fig. 537); second and third instar larva sedentary and
(mM ROUMINS: gn Boo. FO Mego a Bo ERB oes Slory Can cera ae meneaig | a ee eMC T Pen ts ee eee Mantispidae

4'. Prothorax not elongate; forelegs not greatly enlarged or raptorial (Fig. 36); larvae not maggot-like ......... 5

5(4'). Forewings with two or more branches of Rs arising from the apparently fused stems of Rand Rs (Figs. 442-447).

eee dae ts Ace Wo ee eee I Ee aA eon hae Nee oly at lene ees Hemerobiidae
5’. Forewings with all branches of Rs arising from a single sector separate from R, (Fig. 36). ....-......-.. 6
OG) sesniennaemonilitonnun bothisexes: oviposition notprojectines e720 a 2 2 ee eae ee if
6’. Antennae of male pectinate, of female filiform; female with an exserted ovipositor as long as body (Fig. 434)... .

PR er tr re ene ee, eck eae ene la ete Olas cece, Ao Sisk. poe ee <3 Dilaridae

7(6). Humeral crossvein recurved, branched; first r-m crossvein in hind wing longitudinal and sigmoid; large species with

Win S[oeR IG Qed OaySyinoTany((EVER (CSAS) eo rome nS ono ale eat ce omens, Co fe Some cma ae Polystoechotidae
7’. Humeral crossvein simple, not recurved or branched; first r-m crossvein in hind wing variable in shape; smaller spe-
GleSsT (MOS. Ummuwin Ors pam) (E128) rack mmr bananas eran Pens Sey Gee yen eve ewer) ecier cau pusuemts 8

8(7'). Vertex convex; wing venation relatively simple: radial sector of forewings without definitive crossveins (Fig. 660),
Sc and R, coalesced near wing tip, costal crossveins not forked, r-m crossvein of hind wings in the axis of wing; size
small, 6 mm in overall length; paired parapsidial sulci absent on the anterior half of the mesonotum; larvae aquatic,
HSCS OO MRSS ONES PS DONS <6 be a 4 & Ged GG cp us Galo to edeGs Bk Dede OG. CuO 0) cede Gp 0. Dede om owt Sisyridae

8’. Vertex flattened; radial sector in forewings with definitive crossveins (Fig. 28); Sc and R, coalesced or spearate near
wing tip, costal crossveins forked or simple; hind wings with r-m crossvein oblique or transverse; size larger; paired
parapsidial sulci present on the anterior half of the mesonotum; larvae terrestrial, sometimes associated with ant or
WETTING INENES “a. oe beg Ae Ane aka cua awe Gea eet tes rae Serene ORE ae en ORG ners icone iec cnr 9

176 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES
Volume 53, No. 12

9(8’). Wing apices rounded, not falcate; costal crossveins simple, not forked; setae on wing veins shorter than cell widths
and setae on body shorter than abdominal width; no scales on wings, legs, or body; Rs swinging away from Rj,
R,-Rs area broadiand containin -manyscross veins (Eps 2.8) sei cure Chrysopidae

9’. Apical portion of the hind margin of forewings straight or concave (falcate), apex more or less acute (Fig. 27); costal
crossveins forked; wings and body very hairy, especially the hind margin of wings (wing vein setae longer than cell
width and body setae longer than width of abdomen); wings may bear scales along base of longitudinal veins; R,—Rs
atea marrow withifewiCrOSSVEIMS. ase 4o<.5 Geese le me eel oleae ie Be Ste eee Berothidae

FAMILY ASCALAPHIDAE

by
Norman D. Penny

Ascalaphids, or owlflies, are relatively abundant in Costa Rica, and because of their large size and
frequent attraction to light are often seen in collections. This family is quite distinctive. Adults super-
ficially resemble dragonflies, except for their long, clubbed antennae (Fig. 1). Larvae are highly pred-
atory, with one subfamily (Ascalaphinae) feeding on soil surface insects and the other
(Haplogleniinae) feeding on leaf-dwelling insects. Eggs of Ascalaphinae are often laid at the end of
dead twigs with false eggs (repagulae) placed below them (Fig. 4). The repagulae are thought to havea
repealant effect on small predators, such as ants. The young larvae spend the first few days of life
grouped together at the end of their birth twig before dropping to the ground to begin life at the soil sur-
face as lone predators. Larvae of Haplogleniinae have extremely flattened bodies and blend in very
well with the fungus spots often seen on tropical leaves (Fig. 5). They rest quietly on leaves with their
long mandibles wide open for days until an unwary insect comes near. Then, with one quick move-
ment the prey is seized. Their metabolic rate must be very low and they can go for long periods without
feeding. They are particularly fond of mealy-bugs (Pseudococcidae) and other slow-moving,
soft-bodied prey. Adults are most active just at dusk when they dart back and forth after small flying
insects. During the day adults rest on vegetation, the Haplogleniinae with wings open and
Ascalaphinae with wings closed over the body. Adults of the two subfamilies are easily separated be-
cause the Ascalaphinae have a very distinctive partition of the compound eye into upper and lower
halves (Fig. 3). This is lacking in Haplogleniinae (Fig. 2) in which the eyes are hemispherical. Accu-
rate species identification in this family is complicated by relatively uniform male genital armature,
frequent sexual dimorphism, and considerable intraspecific variation in wing patterning.

KEY TO THE GENERA OF COSTA RICAN ASCALAPHIDAE
(modified from Penny 1981)

1. Eyes entire, hemispherical, not divided by a transverse sulcus (Fig.2) ........ Subfamily Haplogleniinae . . . 2
1’. Eyes divided by a transverse sulcus into two equal or unequal parts (Fig.3) ...... Subfamily Ascalaphinae . . .5
2(1). Forewing axillary angle bearing an acute projection at base (Fig. 11) ................ Neohaploglenius
2 Forewing axillary anele‘at baseiotanal maresinmeht-angpledi(Eigs 7) = 9) ee) eee eee 3
3(2)) "Antennae loncenthanidistance tojsecond Rs) forksotsonre wines sei ey eal ite ee 4
355 Antennaeishorter thanidistancetoisecond RSiforksotstore win ss-seee einen er en) eine Ascalobyas
4(3). Hind wing with 2A present, long; forewing with dark coloration along costal margin (Fig. 7)... . . . Haploglenius
4’. Hind wing with 2A absent or extremely short; forewing with translucent clear coloration along costal margin (Fig. 6)
sug inde ee 2! oe eg: are ei RS, See e- 2 ees l P eeeecae eeCey Amoea

5(1'). Hind wing CuP straight (Figs. 12-16); antennae usually as long as forewing (occasionally somewhat shorter)
ee eee cece eee a eter Ones Ch Ceo cel Of om. One Tor oy cede seen of 65 2 Ameropterus

5S’. Hind wing CuP clearly sinuous (Figs. 22-26); antennae much shorter than length of forewing ............ 6

PENNY: LACEWINGS OF COSTA RICA 177

6(5'). Hind wing long and narrow, costal and anal margins approximately parallel (Figs. 22-26) ......... Ululodes
6’. _ Hind wing subtriangular, much wider at mid-length than near wing base (Figs. 17-21). ........ Cordulecerus

SUBFAMILY HAPLOGLENIINAE
Amoea Lefebvre, 1842

Amoea is probably the easiest genus of haplogleniine owlflies to recognize because it is the only
genus with absolutely no dark pigmentation anywhere on either wing, except at the extreme base. All
other genera of haplogleniine owlflies have at least some pigmentation along the costal or subcostal
margin of the wings. Although relatively large owlflies, they are the smallest representatives among
the Costa Rican Haplogleniinae. This genus also has proportionally broad hind wings, which often
protrude dorsally above the forewings when closed. In these respects they closely resemble Ululodes
and some pale-winged forms of Cordulecerus in the Ascalaphinae. However, a look at the undivided
eyes will quickly place these insects in the Haplogleniinae.

Described species of Amoea display almost no differences from each other except slight differ-
ences in size and intensity of a single color pattern. It is likely that many names will be synonymized
in the future, and it is possible that only a single species exists.

Only one nominal species of Amoea is found in Costa Rica, Amoea vacuus.

Amoea vacuus (Gerstaecker, 1893)
(Fig. 6)

DIAGNOSIS. — As mentioned above, all described species of Amoea are very similar, if not iden-
tical. Thus, current identification is based solely on geographical distribution.

GEOGRAPHICAL DISTRIBUTION. — Weele (1908) mentioned this species from Honduras. Fur-
ther specimens have been seen from Mexico (Chiapas State) and Panama. Within Costa Rica, it has
been collected most frequently in the drier northeastern part of the country but has been found in the
south, on both Caribbean and Pacific coasts.

ALTITUDINAL DISTRIBUTION. — Elevation 30m at Lomas Barbudal Biological Station
(Guanacaste) to 1300 m at Las Mellizas in La Amistad National Park (Puntarenas).

ADULT FLIGHT PERIOD. — The vast majority of specimens have been collected in May, June,
and July.

HABITAT PREFERENCE. — Unknown.

Ascalobyas Penny, 1982

This genus of haplogleniine owlflies is characterized by very short antennae, usually reaching
only as far as the origin of the radial sector in the forewing. They appear to be large, forest dwelling
species, frequently attracted to light. Three species are currently known, distributed from Bolivia and
central Brazil north to Mexico, with one record from Texas. Two species are known from Costa Rica.

KEY TO COSTA RICAN ASCALOBYAS

IEP Anexaoigoncwinoypalespterost pmardanka (Hig 00) eee meee ee oes ye ee ee 4. microcerus
apexonbothwines dark- pterosiiomaltareaipale (Eig 8) 15 4566-46 500586 oe ee ae ee 4. albistigma

178 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES
Volume 53, No. 12

Ascalobyas albistigma (Walker, 1853)
(Fig. 8)

DIAGNOSIS. — The short antennae, pale stigmal region, and darkly pigmented apical fifth of both
wings are distinctive. One other Costa Rican species, Haploglenius luteus, also has a pale stigmal area
and darkened apex of both pairs of wings, but the antennae are longer, reaching as far as the first radial
sector fork. Additionally, A. a/bistigma has a tuft of white setae at the base of each wing and the pleural
region of the thorax darkened. Haploglenius luteus has dark setae at the wing bases and a diagonal
pale stripe on the pleural region. Ascalobyas albistigma exhibits sexual dimorphism, with males being
smaller and with dark pigmentation fading to transparency at the apex of both wings. Females have
darkly marked wings up to the wing apex.

GEOGRAPHICAL DISTRIBUTION. — Weele (1908) mentioned this species from Honduras, Pan-
ama, and Colombia; Penny (1981) from the Brazilian Amazon Region and Vogtsberger (1990) re-
ported one specimen from U.S.A. (Texas). Also known from southern Mexico and Belize
(unpublished records). Within Costa Rica it has been collected at GUANACASTE: Barra Honda Na-
tional Park; the west side of Orosi Vulcano at Estacion Maritza; Santa Rosa National Park; LIMON:
Tortuguero National Park; and PUNTARENAS: Carara Biological Reserve. It appears to be fairly
common in Santa Rosa National Park.

ALTITUDINAL DISTRIBUTION. — 300-600 m.

ADULT FLIGHT PERIOD. — May and June, except one collection from Carara Biological Reserve
in April.

HABITAT PREFERENCE. — Unknown.

Ascalobyas microcerus Rambur, 1842
(Fig. 10)

DIAGNOSIS. — The short antennae, dark pterostigmal spot, and pale forewing apex separate this
from all other species.

GEOGRAPHICAL DISTRIBUTION. — From Mexico (Nayarit, Durango, and San Luis Potosi states)
south to central coastal Brazil and Ecuador. In Costa Rica found only at ALAJUELA: San Ramon For-
est Reserve.

ALTITUDINAL DISTRIBUTION. — 900 m.

ADULT FLIGHT PERIOD. — April to June.

HABITAT PREFERENCE. — Unknown.

Haploglenius Burmeister, 1839

With seven Neotropical species distributed from Honduras south to Argentina, Haploglenius is
the most speciose genus of New World haploglentine owlflies. Species of this genus have relatively
long antennae (as long as the forewing to second fork of Rs), forewing with no acute axillary angle at
base, and a dark band along the costal margin of both wings (absent in teneral specimens). They are
found in forested situations where they rest with wings partially open. Several species are attracted to
light. Males have a peculiar flap at the anterior margin of the pronotum which can be flipped back to
reveal a bright white patch. Eisner and Adams (1975) thought this to have a startle function towards
predators. One additional species was described from Papua New Guinea, but this form almost cer-
tainly belongs in another genus. Two species are known from Costa Rica.

PENNY: LACEWINGS OF COSTA RICA 79

KEY TO COSTA RICAN H4APLOGLENIUS

lpeApicaleiithiomhindawinesidarkes(hio* [ian UPei. eee ee ei me So es Be eR ee H. luteus
Manincnwvinesdancenedonlyalongiapicalimaroim(Pis.9)) 2 oa. wn te we ee ee ee ee H. peruvianus

Haploglenius luteus Walker, 1853
(Fig. 7)

DIAGNOSIS. — Superficially this species appears very similar to Ascalobyas albistigma, but the
longer antennae extended as far as the forewing second radial sector fork, dark setae at the wing bases
and diagonal pale pleural stripe are all characteristic of Haploglenius.

GEOGRAPHICAL DISTRIBUTION. — Honduras south to Bolivia and Peru according to Weele
(1908). Additional specimens from southern Mexico (unpublished data). In Costa Rica collected at
HEREDIA: Braulio-Carrillo National Park, Est. Magsasay; LIMON: Cerro Cocori; and Hitoy-Cerere
Biological Reserve; PUNTARENAS: Osa Peninsula, Rancho Quemado; Carara Biological Reserve.

ALTITUDINAL DISTRIBUTION. — 50-300 m.

ADULT FLIGHT PERIOD. — March to May, and September (possibly two generations).

HABITAT PREFERENCE. — Unknown.

Haploglenius peruvianus Weele, 1908
(Fig. 9)

DIAGNOSIS: The lack of a prominent axillary angle in the forewing, antennae as long as length of
forewing to second radial sector fork, and transparent fore- and hind wings characterize this species.
The only other species of this genus in the country is H. /uteus, which has a prominent dark apex to the
hind wing. Haploglenius peruvianus closely resembles H. costatus from southern South America,
which has a uniformly darkened costal area, including pterostigma. The pale stigmal area of H.
peruvianus 1s distinctly set off from the darkened costal margin.

GEOGRAPHICAL DISTRIBUTION. — Costa Rica and Panama south to Bolivia, Argentina
(Misiones Prov.) and Peru. In Costa Rica collected at HEREDIA: 2 km south of Pueblo Nuevo
Sarapiqui; PUNTARENAS: Carara Biological Reserve; and LIMON: Hitoy-Cerere Biological Re-
serve.

ALTITUDINAL DISTRIBUTION. — Up to 100 m elevation.

ADULT FLIGHT PERIOD. — April, July, and September.

HABITAT PREFERENCE. — Unknown.

Neohaploglenius Penny, 1982

This genus was erected for two species from Central America: Haploglenius angulatus
Gerstaecker and Haploglenius flavicornis McLachlan. They appear similar to Haploglenius with a
dark costal wing margin and long antennae. However, the wings are narrower with associated
venational loss, and there is a distinctive axillary angle projection at the base of the forewing, which is
absent in Haploglenius. The two described species are very similar and are herein considered syn-
onyms. Neohaploglenius rondonianus Penny has subsequently been described from the southwestern
Brazilian Amazon.

180 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES
Volume 53, No. 12

Neohaploglenius flavicornis (McLachlan, 1871)
(Figs. 2, 11)

DIAGNOSIS. — This is the only species of haplogleniine in Costa Rica with the distinctive elon-
gate projection at the base of the forewing (axillary angle) .

GEOGRAPHICAL DISTRIBUTION. — Records indicate a distribution from Mexico (Hidalgo and
Sinaloa States) south to Panama. This is probably the most commonly collected haplogleniine
ascalaphid in Costa Rica and is in all likelihood found throughout the country at elevations under

1500 m. In Costa Rica collected at the following localities: ALAJUELA: San Ramon Forest Reserve;
2 km southwest of Dos Rios; CARTAGO: Tuisito 1200; GUANACASTE: Guanacaste National
Park; Rodriguez National Wildlife Refuge; LIMON: Hitoy-Cerere Biological Station
PUNTARENAS: Monteverde Biological Reserve; 14.1 misoutheast of Esparta; Osa Peninsula; SAN
JOSE: Escazu; Finca La Caja; Patarro-Guatuso.

ALTITUDINAL DISTRIBUTION. — 10—1520 m.

ADULT FLIGHT PERIOD. — March to October.

HABITAT PREFERENCE. — Collected in narrow tree line between open pastures near Monteverde
Biological Reserve.

REMARKS. — Haploglenius angulatus Gerstaecker, 1894 is anew synonym of Neohaploglenius
flavicornis. There can be considerable developmental variation in wing pattern. Approximately
one-quarter of specimens of both sexes have smoky dark tints to the wing membrane, perhaps due to
killing reagents. Upon emergence, as wings dry and dark patterns develop, the dark subcostal pattern
appears before the costal pattern, so that some recently emerged specimens have wings with darkened
subcostal area, but translucent costal area. Likewise, the antennae appear yellowish upon emergence,
but darken with age.

SUBFAMILY ASCALAPHINAE
Ameropterus Esben-Petersen, 1922

This genus is most easily characterized by the hind wing venation, which includes the MA, MP,
and Cu forming a distinctive set of three straight parallel veins angled diagonally across most of the
hind wing (Figs. 12-16). The other two genera of Costa Rican Ascalaphinae, Cordulecerus and
Ululodes, both have Cu with an arc at mid-length, thus not parallel to the medial veins (Figs. 17—26).
There has been some confusion as to the validity of this genus because one North American species,
Ameropterus excisus Hagen, has turned out to be the male of Ululodes quadrimaculata (Say). The
deeply incised, hind wing, anal margin of the male, reduces the wing width at a point where Ululodes
normally has the arched Cu curvature. The narrowed wing base and parallel MA, MP, and Cu proba-
bly is a secondary development for this one species. Most species of Ameropterus have antennae as
long as the length of the forewing to the pterostigma, and often are longer than the wings. Most species
of Ululodes have shorter antennae which do not reach to the forewing pterostigma. Male terminalia in
the Neotropical members of this subfamily are very similar, if not identical, and thus not useful in
identification. Ameropterus is an exclusively Neotropical genus, with 21 currently recognized species
ranging from southern Mexico (Chiapas state) to northern Argentina. Five of these species are known
from Costa Rica.

PENNY: LACEWINGS OF COSTA RICA 181

KEY TO COSTA RICAN AMEROPTERUS

1. Small species (forewing length 20 mm or less); hind wing with only six apical forks of CuA; hind wing only about

PEMene iano tone win Oa (HIP el) Per Sarees sR pee ee es bch Je a ete 2 oa A. breviantennis
1’. Larger species (length 25 mm or more); hind wing with 10 or more apical forks of CuA; hind wing nearly as long as
Ace tT Oa (D1 SSeS LG) Sree Careers ee Cer Lan suis s, oee ers cess Me yes Hoy oS a Boe, tee Ve cl <a Gn eG) Om egy 2

2(1'). Hind wing much narrower than forewing; anal area of hind wing with distinct axillary angle (Fig. 15) . A. subripiens
2'. Hind wing similar in shape to forewing; anal area of hind wing with or without an axillary angle, but never as a dis-

inkerett (olor (LATS. A IS) r eos Guay Rene hc cut R OME cn OMe Ee CO ROn! ine nn here be fa, crit Came e Mnry fetes CM Seete ORE WORRIES I 3
3(2'). Pterostigmal area of forewing pale, associated veins widely spaced, not swollen (Fig. 14) ...... . A. mexicanus
3’. Pterostigmal area of wings darkened, associated veins closely spaced, often swollen (Fig. 16) ........... 4
4(3'). Pterostigmal area large, encompassing three or four cells (Fig. 16); pleural area of thorax pale... . . . . A. trivialis
4’. Pterostigmal area small, encompassing two cells (Fig. 13); pleural area of thorax dark... 2... 2... 2. A. consors

Ameropterus breviantennis Penny, 1982
(Fig: 12)

DIAGNOSIS. — This species is readily recognized by its small size and very short hind wings. It
also appears to have a proportionately larger antennal club for the size of its antennae.

GEOGRAPHICAL DISTRIBUTION. — Originally described from the eastern part of the Brazilian
Amazon region and has been subsequently collected in Mexico (Chiapas) and Panama. In Costa Rica
this species has been collected at ALAJUELA: 2 km southwest of Dos Rios; GUANACASTE: 7 km
south of Santa Cecilia; HEREDIA: Magsasay Biological Station, Braulio-Carrillo National Park;
Chilamate; La Selva Biological Station; LIMON: Cerra Cocori; and PUNTARENAS: Osa Peninsula.
This species appears to be most abundant at Pitilla Biological Station in Guanacaste Province.

ALTITUDINAL DISTRIBUTION. — 150—700 m.

ADULT FLIGHT PERIOD. — May to July.

HABITAT PREFERENCE. — Unknown.

Ameropterus consors (Gerstaecker, 1893)
(Fig. 13)

DIAGNOSIS. — This rather large species is characterized by fore- and hind wings of comparable
length; no lobe at the base of the anal margin of fore- and hind wings; and small, dark, forewing
pterostigmata that encompass only two (occasionally three) cells.

GEOGRAPHICAL DISTRIBUTION. — Originally described from Panama, this species has been col-
lected in Costa Rica seven times at four different sites: ALAJUELA, San Ramon Forest Reserve, Rio
San Lorencito; Finca San Gabriel, 2 km southeast of Dos Rios; GUANACASTE, Pitilla Biological
Station, 9 km south of Santa Cecilia; and PUNTARENAS, La Amistad National Park, Las Mellizas
Biological Station, Finca Cafrosa.

ALTITUDINAL DISTRIBUTION. — 600-1300 m.

ADULT FLIGHT PERIOD. — May, with one October record.

HABITAT PREFERENCE. — Unknown.

182 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES
Volume 53, No. 12

Ameropterus mexicanus (Weele, 1908)
(Fig. 14)

DIAGNOSIS. — The pale pterostigma, equally-long fore- and hind wings, and lack of a distinctive
anal lobe at the base of the forewing serve to characterize this species.

GEOGRAPHICAL DISTRIBUTION. — This species was originally described from Mexico and Hon-
duras. It has been collected in Costa Rica from nine specimens at five different locations:
ALAJUELA, San Ramon Forest Reserve, Rio San Lorencito, 5 km north of Colonel Palmarena;
GUANACASTE, Pitilla Biological Station, 9 km south of Santa Cecilia; PUNTARENAS, La
Amistad National Park, Las Alturas Biological Station, Coto Brus; La Amistad National Park, Las
Mellizas Biological Station; and SAN JOSE, Carara Biological Reserve, Bijagual Biological Station.

ALTITUDINAL DISTRIBUTION. — 500-1500 m.

ADULT FLIGHT PERIOD. — Apparently bimodal: May—June, and September—November.

HABITAT PREFERENCE. — Unknown.

NOTE. — The similarity in appearance and collecting sites and dates between A. consors and A.
mexicanus leads one to question the separation of these two species. Because the pterostigma is so
strikingly contrasted in the two forms, they are maintained as separate species for the present, al-
though this is normally a very weak character to use for species separation and these forms may later
be synonymized.

Ameropterus subripiens (Walker, 1853)
(Fig. 15)

DIAGNOSIS. — The distinctively narrowed hind wings of this species immediately separate it
from all other Ameropterus known from Costa Rica. In the forewings, the base of the anal margin has a
small lobe. In the hind wings, the base is very constricted with a very pronounced basal lobe.

GEOGRAPHICAL DISTRIBUTION. — This species was described from Venezuela. There is a speci-
men in the Natural History Museum (London) from Panama. In Costa Rica, this species has been col-
lected only three times, at two localites: HEREDIA, Braulio-Carrillo National Park, Magasay
Biological Station; and PUNTARENAS, Manuel Antonio National Park, Quepos.

ALTITUDINAL DISTRIBUTION. — 80-200 m.

ADULT FLIGHT PERIOD. — February to May.

HABITAT PREFERENCE. — Unknown.

Ameropterus trivialis (Gerstaecker, 1888)
(Fig. 16)

DIAGNOSIS: The combination of uniformly long fore- and hind wings, which lack an anal lobe
and dark pterostigmata that encompass four or five costal cells are diagnostic.

GEOGRAPHICAL DISTRIBUTION. — This species was originally described from Panama. There
are specimens from Mexico (Chiapas) in the collections of the Canadian National Collection and the
University of Nebraska and a specimen from Guatemala in the collection of California Academy of
Sciences. Banks mentioned this species from Costa Rica as early as 1914. This is the most common
species of Ameropterus in the country and has been collected numerous times at the following locali-
ties: ALAJUELA, 20 km south of Upala; Finca San Gabriel, 2 km southwest of Dos Rios; San Ramon
Forest Reserve, Rio San Lorencito; CARTAGO, Juan Vifas; Turrialba; GUANACASTE, Maritza
Biological Station, west side of Orosi Volcano; HEREDIA, Braulio-Carrillo National Park, Magasay
Biological Station; LIMON, Cerro Cocori Sector, Finca de E. Rojas; Braulio-Carrillo National Park,
Quebrada Gonzalez; and PUNTARENAS, Carara Biological Reserve, Quebrada Bonita Biological

PENNY: LACEWINGS OF COSTA RICA 183

Station; Manuel Antonio National Park, Quepos; Osa Peninsula, Rancho Quemado; Osa Peninsula,
2.9 km (1.8 mi) west of Rincon; Golfito; Las Cruces, near San Vito.

ALTITUDINAL DISTRIBUTION. — Sea level to 600 m.

ADULT FLIGHT PERIOD. — February to July, with a peak in May.

HABITAT PREFERENCE. — Unknown.

Cordulecerus Rambur, 1842

Most species of this genus have distinctive pigmentation patterns on the wings. However, unlike
the brightly colored diurnal European Libelloides Schaffer, species of Cordulecerus are crepuscular
and nocturnal. In their most developed form, the wings (particularly the hind wings) are also very dis-
tinctively broadened at mid-length. Identification has at times been confused by the highly dimorphic
wing color patterns of some species. Species of this genus also form mixed-sex aggregations of up to
50 individuals clustered at the tips of tree branches and vines, often overhanging water (Covell 1989;
Hogue and Penny 1989; Gomes-Filho 2000). This behavioral trait is unknown in other ascalaphids.
Cordulecerus can be characterized by the mid-length arch in hind wing vein Cu, and the hind wings
broader than the forewings. This genus is exclusively Neotropical, with 10 species and one subspecies
known from southern Mexico south to central Brazil. Three species are known from Costa Rica.

KEY TO COSTA RICAN CORDULECERUS

1. Smaller species (forewing length 20-28 mm) with series of four or more broken or interconnected dark markings on

Hosteriommind wane sano (Eos 21s. leet a Ra ere ECM es i toa yk ge eye eine C. subiratus
1’. Larger species (forewing length 25-30 mm) with clouded regions or three spots on posterior hind wing margin

(Fis, DO) ob) gh pace de sine sie ote RES ols gaat eect aaa iniesmtaitier it eter here nA ir Sek er anw eS |S 2
Pps Nomarksspotsionhind wine ibasalihalfi (Rigs: L7819)) yess. Ree ei): SOS A). ee 3)
Pee arkeareasson hind wine basalihalt (Rigs lS 20)\t202 ai a es Res @ ek se ee eerie 4
3(2). Apex of fore and hind wings clear and transparent or somewhat evenly enbrowned (Fig. 17).. . . male C. inquinatus
3’. Apex of fore and hind wings contrastingly darker tinted than base (Fig. 19)... ......... male C. praecellens

4(2'). Base of hind wing more or less evenly translucent, with three dark spots along hind margin (Fig.20) ........
etn, a ee ee ed ak ee. ee Se Pe ee Gee RID See Peo female C. praecellens
4’. Base of hind wing dark, with dark area along most of hind margin (Fig. 18) .......... female C. inquinatus

Cordulecerus inquinatus Gerstaecker, 1888
(Figs. 17, 18)

DIAGNOSIS. — This species is highly dimorphic, with wings uniformly clouded with pale brown
(males), or dark areas along the costal margin of the forewing and over much of the hind wing (fe-
males). Hind wings have a broad basal expansion along the posterior margin. The location of pigmen-
tation on the female wings is similar to that of C. subiratus although more diffuse and less fragmented,
but the pattern of males is very different (no discrete markings), and C. subiratus is a much smaller
species.

GEOGRAPHICAL DISTRIBUTION. — This species has been reported from Costa Rica, Panama, Ec-
uador and Peru. Within Costa Rica it has been collected repeatedly from: ALAJUELA: San Ramon
Forest Reserve, Rio San Lorencito. It is also known from SAN JOSE: Braulio-Carillo National Park,
Quebrada Gonzalez.

ALTITUDINAL DISTRIBUTION. — 480-980 m.

ADULT FLIGHT PERIOD. — May to July, with most records from May.

184 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES
Volume 53, No. 12

HABITAT PREFERENCE. — Unknown.

Cordulecerus praecellens (Gerstaecker, 1884)
(Figs. 19, 20)

DIAGNOSIS. — Medium-sized species (forewing length 29 to 30 mm). Males have somber wings
created by dark margining ofall veins, with more intense darkening near tip of wings. Forewings of fe-
males are translucent, except for diffuse darkening at wing tips, and hind wings have a series of three
dark spots along the posterior margin (including the apical spot).

GEOGRAPHICAL DISTRIBUTION. — This species 1s known from Costa Rica, Panama, Venezuela
and Ecuador. Within Costa Rica records are from: ALAJUELA, two km southwest of Dos Rios, Finca
San Gabriel; HEREDIA, Braulio-Carrillo National Park, Magasay Biological Station; LIMON,
Hitoy-Cerere Biological Reserve, Hitoy-Cerere Biological Station; and PUNTARENAS, La Amistad
National Park, Las Mellizas Biological.

ALTITUDINAL DISTRIBUTION. — 100-1300 m.

ADULT FLIGHT PERIOD. — Costa Rican records suggest a rather narrow adult activity period
from May to June, at the height of the late spring/early summer rains.

HABITAT PREFERENCE. — Records suggest a preference for forested habitats.

Cordulecerus subiratus (Walker, 1853)
(aigs 21)

DIAGNOSIS. — This is the smallest of the three Costa Rican species of Cordulecerus (forewing
length 23 to 27 mm), and the only one with little or no wing pattern sexual dimorphism. Forewings are
basally diffusely darkened and apically translucent, except for a thin, broken line from pterostigma to
wing apex. Hind wings have a unique color pattern which is completely dark in the basal third and
translucent in the apical 2/3, witha series of elongate dark markings inward from the posterior margin,
some of which are connected.

GEOGRAPHICAL DISTRIBUTION. — This species is known from southern Mexico, Guatemala,
Honduras, Costa Rica, and Brazil. The Brazilian population is so isolated that it may represent a sepa-
rate, cryptic species. Within Costa Rica it has only been collected at: GUANACASTE, Santa Rosa
National Park, Santa Rosa Biological Station.

ALTITUDINAL DISTRIBUTION. — 300 m.

ADULT FLIGHT PERIOD. — There appears to be a short adult activity period. All 16 known speci-
mens were taken in May and June.

HABITAT PREFERENCE. — Unknown.

Ululodes Currie in Smith, 1900

Species can be characterized as having divided compound eyes, no anal lobe at the base of the
hind wing, a hind wing CuP vein which is strongly arched at mid-length, and hind wings of similar
width as the forewings. This is the largest and most frequently encountered genus of New World
ascalaphids, with 22 recognized species ranging from southern Canada south to northern Argentina.
Within Costa Rica three species are known.

KEY TO COSTA RICAN ULULODES

|. Pterostigmata of fore and hind wings similar in coloration, either all dark or all pale (Fig. 25) ............ 2

l’.... Pterostigma of forewing palesthat ofthind! wingidark (Bigsw22, 23))) 4.4.00). . «Sibi. ee ee U. bicolor

PENNY: LACEWINGS OF COSTA RICA 185

PU) sPterostiomataottore-and hind wings dark (ie)25526)ls 5 3 2 ess aon ee ele ee ee U. macleayanus
eee Picrostiemata oftore- and hind wings;pale(Bigs24)) 3. - 22 ee ee eee 3 ee ee U. cajennensis

Ululodes bicolor (Banks, 1895)
(Figs, 22.23)

DIAGNOSIS. — Among the three closely-related, Costa Rican species this one is distinguished by
having pale forewing pterostigmata and dark hind wing pterostigmata. Males generally have com-
pletely translucent wings but sometimes have a small substigmal mark on the hind wings, while fe-
males have the apical fifth of the hind wings darkened.

GEOGRAPHICAL DISTRIBUTION. — This species is currently known from the southwestern
United States, Mexico, Honduras, Costa Rica, and Panama. Within Costa Rica it has been collected at:
GUANACASTE, 14 km south of Canfas; Pitilla Biological Station, 7 km south of Santa Cecilia;
HEREDIA, La Selva Biological Station; 2km south of Pueblo Nuevo, Finca Naranjo;
Braulio-Carrillo National Park, Magsasay Biological Station; LIMON, Tortuguero National Park,
Cerro Tortuguero; Hitoy-Cerere Biological Reserve, Hitoy-Cerere Biological Station;
PUNTARENAS, Osa Peninsula, Rancho Quemado; Corcovado National Park, Sirena Biological Sta-
tion; and SAN JOSE, Escazu.

ALTITUDINAL DISTRIBUTION. — Sea level to 500 m.

ADULT FLIGHT PERIOD. — Encountered throughout the year.

HABITAT PREFERENCE. — Records suggest that this species frequents open areas at forest mar-
gins.

Ululodes cajennensis (Fabricius, 1787)
(Fig. 24)

DIAGNOSIS. — This species has pale pterostigmal spots, and no dark pigmentation on the wings
of either sex.

GEOGRAPHICAL DISTRIBUTION. — Very widely distributed throughout Mexico, Guatemala,
Costa Rica, St. Lucia Island, Venezuela, Guyana, Surinam, French Guiana, Ecuador, Peru, Brazil,
Paraguay, and Argentina. In Costa Rica this species is known from: ALAJUELA, Gonzita; Guatuso,
La Garroba; Dos Rios-Upala, Finca La Selva, GUANACASTE, Finca Jenny, 31 km north of Liberia;
14 km south of Canfas; Palo Verde National Park, Palo Verde Biological Station; Finca La Flor;
HEREDIA, 2 km south of Pueblo Nuevo, Finca Naranjo; Braulio-Carrillo National Park, Magsasay
Biological Station; LIMON, Hitoy-Cerere Biological Reserve, Hitoy-Cerere Biological Station;
Gandoza and Manzanillo Forest Reserve, Manzanillo; Cerro Cocori Sector, E. Rojas Finca, Finca La
Lola, near Siquirres; PUNTARENAS, Osa Peninsula, Rancho Quemado; SAN J OSE, Escazti; Carara
Biological Reserve, Bijagual Biological Station.

ALTITUDINAL DISTRIBUTION. — Sea level to 500 m.

ADULT FLIGHT PERIOD. — Adult activity appears to be bimodal: March to June, and September
to January. Peak activity is in March and April in the San José area.

HABITAT PREFERENCE. — This species prefers open fields near forest edges.

Ululodes macleayanus (Guilding, 1825)
(Figs. 1, 33,25, 26)

DIAGNOSIS. — Characterized by having dark pterostigmata on both pairs of wings. Males have
entirely translucent wings, while females often have a large subapical dark spot on the hind wing
along the costal margin near the pterostigma, sometimes extending to the posterior margin as well.

186 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES
Volume 53, No. 12

GEOGRAPHICAL DISTRIBUTION. — This is the most common and widely distributed ascalaphid
in the New World, and is the most commonly encountered species in Costa Rica, especially in the
northern part of the country. It is known from the southeastern United States, Cuba, Haiti, St. Lucia Is-
land, St. Vincent Island, Costa Rica, Colombia, Venezuela, Brazil, and Argentina. In Costa Rica it has
been collected at: CARTAGO, Turrialba;, GUANACASTE, 14 km south of Canfas; Guanacaste Na-
tional Park, Santa Rosa Biological Station; 30 km north of Liberia, Finca Jenny; 8 km southwest of
Cuajiniquil, Murcielago Biological Station; Los Almendros; Palo Verde National Park, Palo Verde
Biological Station; Rafael Lucas Rodriguez National Wildlife Refuge, Palo Verde; 12 km southeast
of La Cruz, Cerro El Hacha; Barra Honda National Park, 3 km northwest of Nacaume; HEREDIA,
Braulio-Carrillo National Park, Magsasay Biological Station; 2 km south of Pueblo Nuevo, Finca
Naranjo; and LIMON, Cerro Cocori Sector, E. Rojas Finca.

ALTITUDINAL DISTRIBUTION. — Sea level to 300 m.

ADULT FLIGHT PERIOD. — Encountered year round.

HABITAT PREFERENCES. — This species, like others in this genus, appears to prefer dry, open or
scrub areas near forest edges. Often these areas include cultivated fields.

FAMILY BEROTHIDAE

by
Norman D. Penny

Berothids, beaded lacewings, are fairly common in the warmer parts of North America where ter-
mites are abundant. Some species of Lomamyia are known to live in termite nests as larvae, feeding on
the termites by injecting a paralyzing toxin into the termite, then sucking out body fluids. Termites ap-
parently never recognize these larval berothids as nest intruders. In South America, berothids are
much rarer, and most austral species belong to a separate, primitive subfamily. Central American
berothids were first described by Adams (1990). Only one species occurs in Costa Rica, Spiroberotha
sanctarosae.

Spiroberotha sanctarosae Adams, 1990
(Fig. 27)

DIAGNOSIS. — Like most berothids, this species superficially resembles a brown lacewing in
size and coloration, but is more delicate in appearance, with narrower falcate wings and long, abun-
dant pilosity.

GEOGRAPHICAL DISTRIBUTION. — This species is only known from Costa Rica and appears to
be confined to GUANACASTE, in Santa Rosa National Park; south of Canfas; and at La Pacifica, just
north of Canas.

ALTITUDINAL DISTRIBUTION. — Near sea level.

ADULT FLIGHT PERIOD. — December to May, indicating adult activity during the driest time of
the year in Guanacaste Province.

HABITAT PREFERENCE. — Known hosts for New World berothids include only dry-wood ter-
mites, primarily Reticulotermes spp. Thus, the open, semiarid, deciduous woodlands of Guanacaste
Province seem ideally suited for both termites and berothids.

PENNY: LACEWINGS OF COSTA RICA 187

FAMILY CHRYSOPIDAE

by
Norman D. Penny

The green lacewings constitute the second largest family of the order Neuroptera. However, in
humid tropical forest regions chrysopids far surpass the myrmeleontids in species richness, while
myrmeleontids are more speciose in more arid areas. This general observation holds true for the Costa
Rican fauna, with the Chrysopidae being overall the most abundant and specious family, and with the
Myrmeleontidae reaching their greatest abundance and diversity in the relatively dry Guanacaste
Province.

A normal chrysopid egg is oval and placed at the end of a long filamentous stalk which raises the
egg above the substrate. Some species place their eggs singly, others in large groups, and some even
intertwine the stalks to form a mass of eggs. Female green lacewings are able to sense chemicals in the
air emitted from honeydew and other concentrated secretions of phytophagous insects. Eggs are then
laid close to these initial food sources. Attempts have been made to enhance the number of eggs laid in
agricultural areas by spraying articifical “honeydew” onto crops to attract female chrysopids for
oviposition. These insects are among our most beneficial insects, because larvae feed on eggs, larvae
and adults of a wide variety of phytophagous homopterans and other relatively sedentary insects.
Over the approximately two to three week period it takes for the larva to develop and pupate, hundreds
of small insects or thousands of insect eggs can be consumed. Some green lacewing larvae have tho-
racic projections and long setae to hold the dried skeletons of prey and other bits of debris over their
bodies so that the larva is well protected from visual predators, as well as from other predators and par-
asites. This type of larva is called a “trash-carrier,” and in the forest they appear as small pieces of de-
bris slowly moving along a leaf, twig or tree trunk. Adults feed on pest insects, honeydew, and pollen.
Adults of the genus Chrysoperla, which are ubiquitous in agricultural areas, are known to feed on
grass pollen (Ken Hagen, personal communication).

The most recent generic revision of the Chrysopidae (Brooks and Barnard 1991) lists three
subfamilies, two of which are found in Costa Rica. Only two species of the Apochrysinae are currently
known from the country, while three tribes and 72 species of Chrysopinae are found here. The
chrysopid fauna of Costa Rica is very rich, especially for such a small country, and more species can
be expected to be encountered in the future.

The genera of Chrysopini are very difficult to identify to genus based solely on females. The gen-
era of this tribe are recognized mostly by elements of the male genitalia. Thus, the generic key uses
non-genitalic characters for other tribes, but relies on male genitalic structures within the Chrysopini.

KEY TO THE GENERA OF COSTA RICAN CHRYSOPIDAE
(modified from Henry et al. 1992)

1. Basal antennal segments wider than long; rather stout, wide-bodied species (Fig. 133); male with tignum absent,
gonapsis may be present as two separate projections or a two-horned plate; female with or without praegenitale
eee ae. 5s Tinie Helavapenvemim ears wae aor ft. SP Mek eae a se boat 4 wap hos, i we NGCOFIMG

IM. Basal antennal segments longer than wide; rather thin, slender-bodied species (Fig. 35); male with tignum present
or absent (Fig. 29); gonapsis, if present, not a two-horned plate; female without praegenitale .......... 2

2(1') No dark spot at base of pterostigma (Fig. 141); antennae shorter than 1.3 times forewing length; male with tignum
present or absent, gonapsis present or absent (males required for further identification in this key)
Spey Atak ees Ee As U® DEY SO MMer PETIA ad hy See Sey ots Peek ty he, EBM Gaede tage Ps eee es WY Ge ea ant Le een
hig A dark spot at base of stigma (Fig. 48); antennae distinctly longer than 1.3 times forewing length; male without
(MeO MOM Gs o ole Gedo a Bee lala anne eraeencmems meio. cue Eom en ENE Roe COREE temic ne cro aut rom 3

188 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES
Volume 53, No. 12

3(2'). In forewing, basal subcostal crossvein and intramedian cell absent; distinct blister-like bullar spots present (Fig. 33).

BRO, ocr Subfamily Apochrysinaé «os e.< ik cet eee yn eh wy ge ee oS
Bie In forewing, basal subcostal crossvein and intramedian cell present; bullar spots absent (Fig.45) .........

Butea et Tribe Leucochrysimit .0 265 2 6 ce ee ke so es eS ey eee op © oot eee
AQ): Male without tignuml wy. 42. cr Soe Goel Sa A el Sls Goes S21, 7
4’. Male! with:tignuimi (Big: 29)in 03.2) & aeo ei af le? te te Se es a) ge Bl 5
5(4'). Male with pseudopenisi(Fig:, 29)... onus ws Si) © as ete Sarat tli chad 212 eek a en 6
Sie Male:without pseudopemis:;.. | secs. ca 6 a, 8 pgs oes eS sie cue oe ee Chrysoperla

6(5). Male with enlarged or modified antennal bases and/or with facial horns (Figs. 314, 315); gonapsis (Fig. 29) usually
present £sa¥ yo i Peery eee ae hs ee es Se ee Soi i Meleoma
6’. Male without modified antennal bases or facial horns (Fig. 317, 320); gonapsis absent ........ Plesiochrysa

7(4). Male with base of ectoproct not extended basally to articulate with base of sternite 8 + 9; ectoprocts fused

dorso-medially at base; dorsal apodeme oriented vertically through callus cerci (Fig. 325).......... Ungla
Ms Male with base of ectoproct extended basally to articulate with base of sternite 8 + 9; ectoprocts not fused

dorso-medially; dorsal apodeme oriented horizontally, often branched around callus cerci and ventral branch ex-

tended) beneath ectoproctasiapicall pomt|(ig 2162) 5 ep 2 eae ee eee 8

8(7'). Male with two horn-like structures on gonarcus or arcessus (Fig. 169); gonapsis elongate (Fig. 170) Ceraeochrysa

8’. Male without horn-like structures on gonarcus-arcessus; gonapsis absent or present, but not elongate (Fig. 281) 9
9(8)).- ~Malenwith pseudopenis(Figs29)\\5 2. SSI 28 Bs 2, Ga) 2 RSE LY) Se Meleoma
oF Male without pseudopenis, cucrs yess a fee ee cen wpe ass Mec Ge eee ee 10

10(9'). Mandibles blunt-tipped; wings usually with narrow costal area; male without gonapsis Chrysopodes (Neosuarius)

10’. Mandibles with slender fang-like tips; wings usually with wide costal area; gonapsis present in males of a few
SPECIES Pree NN Ne Ret ee eae Se, ee EE eee EEN Oe ee ome Chrysopodes (Chrysopodes)
11(3’). In forewing one crossvein from MP to R + MA present before origin of PsM (Fig. 36). ....... Berchmansus
Nts In forewing crossveins from MP to R + Ma absent before origin of Psm (Fig. 42) ................ 12
12(11’). Forewing with costal area very broad with sigmoidally undulating crossveins (Fig. 129 ........... Vieira
Ie Forewing wath costal area beanmng stralshticrossvems) (ies 42)) 2 5 ae ee ci eee 13

13(12'). Forewing with intramedian cell triangular, MP, joining MP, before end of cell (Fig. 63) . . Leucochrysa (Nodita)
ye Forewing with intramedian cell quadrangular, MP, not joining MP, (Fig.60) ..........-.....-- 14

14(13). Forewing pterostigma with a large black spot extending back to radial sector (Fig. 42); costal stigmal area usually

twiceras) broadsasisubcostallstiemal areas -) sea 2 es ce ee) cen ee ec Gonzaga
14’. Forewing pterostigmal mark much smaller (Fig. 60); costal stigmal area rarely twice as broad as subcostal stigmal

ALA BAL 3 hes Bae ee ee COS Ek Ce Maye de Ree Pat MNase ne ees Leucochrysa (Leucochrysa)
5G) Forewins withvanlarcion seven or more) blister-like bullae\(Fie533)) eee) fae) ene Loyola
IS Rorewing wath onlyitwolblisterlike) bulllaey(Eio505 (0) es eear nce neie rrr ee Domenechus

SUBFAMILY APOCHRYSINAE

These are the largest and broadest winged of green lacewings. Their flight through the forest is
slow and graceful, much like that of a translucent-winged butterfly. Little is known of their biology.
They are usually found resting under leaves well within the forest. Their broad wings do not allow
them to fold the wings roof-like over the abdomen, as in other chrysopids, but rather they are held hori-
zontally when at rest. There are only 26 species and subspecies in 12 genera worldwide (Brooks and
Barnard 1990), and they are seldom collected. In the New World there are four genera and seven spe-
cies, distributed from Mexico and the Dominican Republic south to southern Brazil. All New World

PENNY: LACEWINGS OF COSTA RICA 189

species have raised, shiny, dark spots called bullae on the forewings. Two genera and species have
been collected in Costa Rica.

Domenechus Navas, 1913

This genus is easily recognized by the two dark bullae on the forewing and the lack of bullae on
the hind wing. There are two described species in this genus: Domenechus marionella
(Guerin-Ménéville) from the Amazon Basin of Brazil and Domenechus mirifica (Gerstaecker) from
Central America.

Domenechus mirifica (Gerstaecker, 1888)
(Figs. 20-32)

DIAGNOSIS. — As for the genus, see above.

GEOGRAPHICAL DISTRIBUTION. — Gerstaecker (1888) originally described this species under
the name Apochrysa mirifica from Panama. In 1913 Navas described this same species under another
name, Domenechus sigillatus from Guatemala. Within Costa Rica, this species is known only from:
PUNTARENAS: Golfito, and La Suerte.

ALTITUDINAL DISTRIBUTION. — The Golfito records are from the lowlands near sea level.

ADULT FLIGHT PERIOD. — The specimen from La Suerte was captured in August whereas the
Golfito specimens were collected in July.

HABITAT PREFERENCE. — Other apochrysines have been collected in primary forest and the
Golfito records on the Osa Peninsula also suggest a lowland primary forest habitat.

Loyola Navas, 1913

This genus is immediately recognizable by the numerous ( >3) blistered spots (bullae) on the
forewing, presence of pigmented blistered spots on the hind wing, and the multiple rows of forewing
gradate veins forming a reticulate web of crossveins in the distal half of both wings. There are two rec-
ognized species in this genus, ranging from Mexico to Brazil. Only one, Loyola croesus, has been col-
lected in Costa Rica.

Loyola croesus (Gerstaecker, 1893)
(Figs. 33-35)

DIAGNOSIS. — This species is unmistakable. It is not only the largest green lacewing found in the
New World, but has broader wings than others, and a characteristic are of seven dark bullae across
each forewing.

GEOGRAPHICAL DISTRIBUTION. — This species is known from Costa Rica, Panama, Venezuela,
and eastern Peru. Within Costa Rica it has been collected only twice: CARTAGO, Santa Teresita de
Lajas, near Turrialba; and Tuis.

ALTITUDINAL DISTRIBUTION. — So far, in Costa Rica this species is only known from
mid-elevation levels on the Caribbean side of mountain slopes near Turrialba and Tuis.

ADULT FLIGHT PERIOD. — The only recorded date for Costa Rica is in June.

HABITAT PREFERENCE. — Although there is no information on habitat preferences for this spe-
cies, apochrysines collected in Africa, Australia, Papua New Guinea and the Brazilian Amazon have
all been associated with understory vegetation in primary forests.

190 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES
Volume 53, No. 12

SUBFAMILY CHRYSOPINAE
TRIBE LEUCOCHRYSINI

The leucochrysines are the dominant group of green lacewings in Neotropical forests. They are
characterized by having very long antennae that extend beyond the wing apices, and a pseudomedial
vein in the forewing that curves directly into the outer gradate series of crossveins. The pterostigma is
darkly pigmented, although occasionally, particularly in teneral specimens, this pigmentation is not
developed. The Chrysopini genus Chrysopodes occasionally has a species with some darkening of the
pterostigma. However, in general this characteristic is useful for quick recognition. Larvae are
trash-carriers. Although dominant in forest habitats, they are only infrequently seen in more open hab-
itats, such as cultivated lands.

This is a very large tribe with many undescribed species. One estimate (Adams 1978) puts the to-
tal number of valid species in the subgenus Leucochrysa (Nodita) at 200 or more. There are four gen-
era found in Costa Rica, with Leucochrysa (Nodita) being by far the most speciose and abundant.

Berchmansus Navas, 1913

This is the only leucochrysine genus with a presectoral crossvein. Three species have been de-
scribed, distributed from Costa Rica to central Brazil. Only one, Berchmansus elegans has been col-
lected in Costa Rica.

Berchmansus elegans (Guérin-Ménéville, 1838)
(Figs. 36-38)

DIAGNOSIS. — This is the only species of Berchmansus known from Costa Rica, so characters
that define the genus will also serve to distinguish this species. Additionally, the heavily pigmented
forewing (Fig. 36) is found in no other species of green lacewing from this region, except perhaps
Vieira leschnaulti, which is much larger and has a quite different pigment pattern.

GEOGRAPHICAL DISTRIBUTION. — This species has previously been collected in Surinam and
Brazil. This is the first record for this species from Central America and Costa Rica. It was found at:
PUNTARENAS: Las Alturas, near San Vito in La Amistad National Park.

ALTITUDINAL DISTRIBUTION. — 1500 m.

ADULT FLIGHT PERIOD. — The only record available is in May.

HABITAT PREFERENCE. — Unknown.

Gonzaga Navas, 1913

Gonzaga is closely related to Leucochrysa (Leucochrysa) having in common with this subgenus
the quadrate intramedian cell. Species of Gonzaga usually have a darkly pigmented frons and vertex
of the head and mesonotum of the thorax, while the pronotum is pale green. The wings are usually
more heavily pigmented than in Leucochrysa, especially in the area of the pterostigma, where the dark
pigment will extend from the pterostigma into the radial area. There are seven currently recognized
species in this genus, distributed from Guatemala south to Brazil and Peru. Two species have been
collected in Costa Rica.

PENNY: LACEWINGS OF COSTA RICA 19]

KEY TO COSTA RICAN GONZAGA

ieWatk spot at intersection of Psm and outer gradate veins (Fig. 42)... 6. eee G. torquatus
1’. No dark pigmentation on forewing apical area, except around pterostigma (Fig.39) ............ G. palliceps

Gonzaga palliceps (McLachlan, 1867) [new combination from Leucochrysa|
(Figs. 39-41)

DIAGNOSIS. — This species has the least wing pigmentation of any of the Gonzaga species, with
dark markings confined to the pterostigmal area. The vertex of the head is also mostly pale, with dark
markings confined to the area just behind the antennae. However, the quadrate intramedian cell, pale
pronotum and dark mesonotum along with dark wing markings in the pterostigmal area, which extend
into the radial area, will distinguish this species as a Gonzaga.

GEOGRAPHICAL DISTRIBUTION. — This species is known from Costa Rica, Guyana, and Brazil.
Within Costa Rica it has been collected at: LIMON, in the Cerere River Valley, both at Pandora and
Hitoy-Cerere Biological Station; and PUNTARENAS, Golfito.

ALTITUDINAL DISTRIBUTION. — Lowlands below 200 m on both coasts.

ADULT FLIGHT PERIOD. — March, July, and September.

HABITAT PREFERENCE. — This species was collected on understory saplings in mature forest
near small streams. They were particularly abundant along a small stream on an isolated wooded hill-
side near the headquarters of the Del Monte Banana Plantation in the Valle de Estrella.

Gonzaga torquatus Navas, 1913
(Figs. 42-44)

DIAGNOSIS. — This species can be characterized as having a very large dark mark over the
pterostigmal area, which extends posteriorly beyond R,. There is a second rhegmal spot where the
outer gradates merge with the pseudomedial vein

GEOGRAPHICAL DISTRIBUTION. — This species is known from Guatemala, Costa Rica, Panama,
Venezuela and Brazil. Within Costa Rica this species has been collected at: ALAJUELA, Finca La
Selva, Dos Rios; LIMON, Pandora; PUNTARENAS, Osa Peninsula, 2.9 km (1.8 mi) west of Rincon;
and SAN JOSE, San José.

ALTITUDINAL DISTRIBUTION. — The Pandora and Rincon specimens are from less than 100 m
elevation.

ADULT FLIGHT PERIOD. — Costa Rican records are from March, June, and September.

HABITAT PREFERENCE. — The Pandora specimen was collected together with Gonzaga
palliceps on understory saplings on a forested hillside along a small stream.

Leucochrysa McLachlan, 1868

Leucochrysa isa very large genus with 161 currently recognized species. The genus is found only
in the New World, primarily in the tropics, where it has become the most diverse genus of chrysopids
in the Neotropical Region. Most species are closely associated with forested areas, although they can
sometimes be found in orchards. The genus is usually divided into two subgenera—Leucochrysa
(Leucochrysa) and Leucochrysa (Nodita).

192 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES
Volume 53, No. 12

Leucochrysa (Leucochrysa) McLachlan, 1868

This subgenus is characterized by having a more or less quadrate intramedian cell in the forewing
(Fig. 45). These species are generally larger than Leucochrysa (Nodita), with longer series of gradate
veins. There are 41 known species ranging from the northeastern United States south to Argentina.
Several unpublished synonyms exist in this subgenus. Six species have been collected in Costa Rica.

KEY TO COSTA RICAN LEUCOCHRYSA (LEUCOCHRYSA)

1. Vertex and mesoscutellum with bright white spots when alive, creamy yellow spots when dead (Fig. 59) .......
Sed cee te Spey Sie EE EE ioe) PS, eas TA = Teed ida eae SSPE ate web EMS bee See eae L. (L.) magnifica
I’. Vertex and mesoscutellum green, pale yellow or brown with various markings, but never with white or creamy yellow

Spots (Fig; 50) esi 2 Se Sa 8 ea le 5 ee he es A cee ee rrr 2
2(1'). Forewing with a dark transverse stripe from base of Rs to anal margin (Fig. 48) ............. L. (L.) clara
2’. Forewing in some individuals with dark markings but never with a transverse stripe from base of Rs to anal margin

(eit: o) eee ee eens 1 Oi EO Se CREE eotiee to 6G 0 oo te 3
3(2))s Pronotumiwith’al dark loneitudinallstipeonteachisidel(R1e5 47) eyes oe ee 4
3) Pronotum: wathionlyslichtdarkomotiline laterally (ie sG2)ie. = eye] ee 2) eee 5
4(3). Frons pale yellow; two dark spots on mesonotum (Fig.47) ....................... L. (L.) adamsi
4'. Frons green; dark arc across anterior margin of mesonotum (Fig.56) ................. L. (L.) lestagei
5(6))sMetascutumi dark: metascutellum’ pale’\(Figs'62))— 5 2") 55%) 42 a ek oie eee ee L. (L.) pretiosa
Sees Metascutum™ipalesmetascutellumidarka(HIes53)) acne seer ce cece enero L. (L.) colombia

Leucochrysa (Leucochrysa) adamsi Penny, 2001
(Figs. 45-47)

DIAGNOSIS. — The narrow dark lateral line of the pronotum and two dark spots on the
mesoscutum are unique among Costa Rican Leucochrysa (Leucochrysa) and the dark region in the ra-
dial area of the forewing is found in some Leucochrysa (Nodita), but not in other Leucochrysa
(Leucochrysa).

GEOGRAPHICAL DISTRIBUTION. — Originally described from Cuba, and subsequently cited
from Honduras (under the name L. navasi). This species has been collected only once in Costa Rica:
LIMON, Pandora.

ALTITUDINAL DISTRIBUTION. — 40 m.

ADULT FLIGHT PERIOD. — September.

HABITAT PREFERENCE. — The Pandora specimen was collected together with Gonzaga
palliceps on a forested hillside along a small stream bordered by understory saplings.

Leucochrysa (Leucochrysa) clara (McLachlan, 1867)
(Figs. 48-50)

DIAGNOSIS. — This is one of the larger species of Leucochrysa. It has a distinctive dark marking
on the forewing running transversely from the origin of the radial sector to the base of the posterior
wing margin. No other species of Leucochrysa has such a mark.

GEOGRAPHICAL DISTRIBUTION. — This species has been recorded from Costa Rica, Panama,
Colombia, Ecuador and Brazil. Within Costa Rica, records are from: ALAJUELA, 20 km south of
Upala; San Ramon Forest Reserve, Rio San Lorencito; GUANACASTE, 14 km south of Canfas; and
PUNTARENAS, San Vito.

PENNY: LACEWINGS OF COSTA RICA 193

ALTITUDINAL DISTRIBUTION. — Pacific Coast lowlands.

ADULT FLIGHT PERIOD. — Numerous records from near Upala indicate the presence of adults
throughout the year.

HABITAT PREFERENCE. — Unknown.

Leucochrysa (Leucochrysa) colombia (Banks, 1910)
(Figs. 51-53)

DIAGNOSIS. — This species is very similar to Leucochrysa pretiosa, except that Leucochrysa co-
lombia has lateral reddish brown markings on the pronotum, with variable amounts of dark pigmenta-
tion on the mesonotum, from an anterior arc to the whole notal area, metascutum pale at least laterally,
and metascutellum dark. The outer gradate crossveins of the forewing are dark, but not margined, and
apical forks at the wing apex are pale.

GEOGRAPHICAL DISTRIBUTION. — This species is only known from Costa Rica, Colombia, and
Ecuador. The only locality within Costa Rica where it has been collected is: PUNTARENAS, La
Amistad National Park, Las Alturas.

ALTITUDINAL DISTRIBUTION. — 1500 m. This may be a mid-elevational species.

ADULT FLIGHT PERIOD. — Mid-February to early March. This is the driest time of the year in the
Talmanca Mountains rain shadow area of La Amistad National Park.

HABITAT PREFERENCE. — Collected on understory saplings, including many Peperonia plants,
in mixed montane primary forest with many large Ficus trees.

Leucochrysa (Leucochrysa) lestagei Navas, 1922
(Figs. 54-56)

DIAGNOSIS: Characteristics include a broad, brown, lateral stripe on the anterior 2/3 of pronotum
and all of the mesonotum; remainder of the thorax is pale green. The frons has a broad transverse red
band.

GEOGRAPHICAL DISTRIBUTION. — This species has been reported from Costa Rica, Ecuador,
and Brazil. Within Costa Rica, records are from: PUNTARENAS, La Amistad National Park, Las
Alturas; Las Cruces (San Vito). Navas (1922) also published two records from the Caribbean low-
lands based on material subsequently destroyed in the Hamburg Museum: LIMON, Rio Reventazon,
Finca Hamburgo; and Las Mercedes, Santa Clara

ALTITUDINAL DISTRIBUTION. — Existing specimens indicate the presence of this species only at
mid-elevations on the Pacific side, or, if Navas’ records are included, also in the Caribbean lowlands.

ADULT FLIGHT PERIOD. — Pacific mid-elevational records are from late February to April, the
driest time of year in this region. One Caribbean lowland record is for August.

HABITAT PREFERENCE. — Collected on understory saplings, including many Peperonia plants,
in mixed montane primary forest with many large Ficus trees.

Leucochrysa (Leucochrysa) magnifica (Banks, 1920)
(Figs. 57-59)

DIAGNOSIS. — This species is recognizable by two, very distinctive spots of bright white, one on
the vertex and the other on the mesoscutellum. These spots in a live individual are easily visible to the
unaided eye. However, they fade to a creamy yellow after death. There is also a diagonal dark band
along the apical forkings of the forewing longitudinal veins.

GEOGRAPHICAL DISTRIBUTION. — Originally described from the southwestern part of the Bra-
zilian Amazon Basin, L. magnifica has also been collected in Costa Rica and French Guiana. Within

194 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES
Volume 53, No. 12

Costa Rica this species has been collected at: ALAJUELA, 20 km south of Upala; and LIMON, the
Estrella Valley south of Limon; at Pandora (also within the Estrella Valley).

ALTITUDINAL DISTRIBUTION. — The Pandora locality is less than 200 m elevation, and the
Upala site is also lowland.

ADULT FLIGHT PERIOD. — Both records from the southern Caribbean coast are from September;
the Upala record is from February.

HABITAT PREFERENCE. — The Pandora collection was made on small understory saplings along
a small stream in primary forest.

Leucochrysa (Leucochrysa) pretiosa Banks, 1910
(Figs. 60-62)

DIAGNOSIS. — This species is very similar to Leucochrysa colombia, except that L. pretiosa has
a completely pale green pronotum, with most of the mesonotum darkened, a dark metascutum and
pale metascutellum. The outer gradate crossveins of the forewing are extensively margined with dark
markings, as are the apical forks at the wing margin, forming a dark transverse apical line.

GEOGRAPHICAL DISTRIBUTION. — This widespread Neotropical species 1s known from Mexico,
Guatemala, Costa Rica, Panama, Colombia, Venezuela, and Ecuador. It is probably the mostly com-
monly collected species of Leucochrysa (Leucochrysa) in Costa Rica, and unlike other members of
the subgenus can occasionally be found in substantial populations. Within Costa Rica this species has
been collected at: ALAJUELA, 20 km south of Upala; 8 km south of Santa Cecilia, Pitilla Biological
Station; CARTAGO, Turrialba; GUANACASTE, Santa Rosa National Park, Rio San Josecito;
HEREDIA, La Selva Biological Station, near Puerto Viejo; LIMON, Rio Reventazon, Finca
Hamburgo; Limon (Ciudad); Talamanca Mountains, Rio Barbilla; Pandora; PUNTARENAS,
Golfito; and SAN JOSE, San José.

ALTITUDINAL DISTRIBUTION. — Sea level to 960 m.

ADULT FLIGHT PERIOD. — Adults have been collected in all months except October and Novem-
ber.

HABITAT PREFERENCE. — This species was collected on understory saplings near a small stream
in mature forest.

Leucochrysa (Nodita) Navas, 1916

This is a very large group of leucochrysines, with representatives found in virtually every tropical
forest in the New World. They are characterized by a triangular intramedian cell in the forewing. Oc-
casionally an individual specimen will have a quadrangular intramedian cell, creating confusion as to
subgeneric placement, but this characteristic is normally stable. This subgenus is sometimes raised to
generic rank. There are 120 recognized species distributed from the southern United States to Argen-
tina, including 24 species recorded from Costa Rica. There is much unrecorded synonymy and many
undescribed species.

PENNY: LACEWINGS OF COSTA RICA 195

KEY TO COSTA RICAN LEUCOCHRYSA (NODITA)

Note: This key uses non-genitalic states, whenever possible, so that both males and females can
be used. Unfortunately, old faded specimens will not key out reliably, so that relatively fresh speci-
mens are needed. Three pair of species in the key will come out together at couplet 6, 9, and 23. They
can only be separated by study of the male genitalia.

Ile Pivelagceiiore wine lengthen Oimm\OrmOnre)i ree ease eset 8 oy et eet se elas fase es eee oe ae py)
PES iZersimallen(torewineilens tl smam\ On less) sean 24 oe a oe em ee he fe ee, cp ey Sa ee 4

2(1). Medial yellow line on thorax and abdomen margined laterally by red; vertex with double red line on both sides

(CEI, PID), pain seshep Ades Giagarchaccla cit yety REO Tore na gstorc to dies rc any eae ec a Ree L. (N.) egregia
Phe Thorax and abdomen without medial yellow line margined laterally with red (Fig. 71); vertex with dark lines, but
OMG OO CRMC STONIDOLIESTCES Hee sem tiie Seeewen city Sete MY orn second a rat oun mee MS 3

3(2'). Vertex with red pentagonal marking; gena red (Fig. 64); four dark spots on mesoscutum (Fig.65) .........

3 Vertex with brown transverse line; gena pale (Fig. 115); mesoscutum without markings (Fig. 116)
re Bi Ga BSA cy CrP UN MRA nmI PRLAEOn Gi hae et te Na FAG a ane nn is eines (elke Lee aia. L. (N.) pallescens

Ue ae NOexecb nitidawan oe anka (Hos UM) ce skater n patateeyls ols Morais st espe a ia lays the ceaproie See Paces aaete-t on on 5
4’. ANTS OH lve | Vomraes OPH TEN Te Gmc cy acl ile dh ued reece mie MRI anes on a eee whens fn cee aoe 10
eH EE Nicixalicava DAL MGarks (RIO IULS) cP cas ee ee ew Es ee eh ee ce ek wl © meee OBE Oe es Gee 6
Ske MPS aGvapalpipalc (IOS) Meee eee Ss Kea nie I Men he en ee nts are Ato tate en ef rete mm ee ays Tames 8
Gis) we asm-bsculcrossvemstallidarklymargmedi(Fis. 17s 525 sess ee ee ee ee ee ee L. (N.) postica
6’. Est = PSGuuGLossviciusmonGarkivemanoine ds (EI C.0/2) |e oc temic ees Gy st Gaye Guns eee 2 eee ese eee 7
i(G))=) Dorsal’horns of male-gonarcus absent; arcessus unipartite <2 2.0... 2... tet ee ee L. (N.) indiga
The Dorsal horns of male gonarcus absent; arcessus tripartite .................++-+--- L. (N.) askanes

8(5'). Base of forewing costal vein dark; transverse reddish brown band below antennal bases interrupted medially
me eae SPE) nde Na ce coe Fe iad Yorn RECO cowl Ge (ae bey & oegsh ie ey ap ota wees Lek ners L. (N.) antica

8". Base of forewing costal vein pale; transverse reddish brown band below antennal bases complete ........ 9
Ose) yee OHOCOLMUA ol eonatcalicomplexmiattened, broad. 2) 522s. .5s 2.5555) eee | L. (N.) cornesta
2). Gonocornua of gonarcal complex elongate, taperedto point ..................... L. (N.) lenora
NO Ce) ae ronommmunitonnly pale(Ric, IZS)R at See ee es eee es ts Seco es oe o 11
10’. Rronotumnvaniablyamarkedawithired ion browii(EiGa86)s ace os cial-me) ce see keno) oartt-t sueuee cen cae seen 12

11(10). Frons with transverse dark or red band (Fig. 127); gonarcus variously shaped, but without double dorsal lobes
ee ee en er eee ENS, SR er) She eS Nie Git) See. ee eo ss eae yeas. L. (N.) virginiae

11’. Frons white, without transverse dark band (Fig. 124); gonarcus bearing double blunt dorsal lobes in addition to
ALGessusTAnG Sharp lyspointed CntOPLOCESSES nara eat, ee ee) eee eee eke Gu ap teeta L. (N.) serrei
MO aMaxilanyapalpiipale tis ws er yk. era i cua EA isan ac, “ME, PL ae eed, 17
12%. NMaxalllanvapal pit ankcarraty te: Meer can esac Vad Posie eogborord aad to a.e-are iS cae ee 13
IB Gea aviesonotumnwithstwodarkmedialispots'.. seas 2 a cee an en eo A ee ee te eee 14
132 Mesonotum completely pale medially, or with thin transverse line across anterior margin ........... 15
12S) Skorewinewlencth—<lSmamaclypeusypdle oui eha-wcne ages Stes) 2) eee) sey es See ee cis L. (N.) caucella

14’, ROLE Win culenothie-ali/ammachypeusidatke fem, Gehan cusee © 3 eee a On Sy ye, Ae i tae ees L. (N.) superior

196 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES
Volume 53, No. 12

15(13’). Complete transverse dark red band across frons beneath antennal bases; scape with ventral longitudinal dark stripe
Soke yeh oh cob iss cicteppns clghe pdt SoalS 2M Boy tag Sa 1s SC Sede a Re ee a L. (N.) duarte

ISMs Dark marks on frons confined to lateral areas near gena; scape completely pale ventrally ............ 16

16(15’). Vertex completely pale; mesoscutum pale; mesoscutellum with dark lateral spots; scape dorsally with one stripe;

pronotim wathoutmed@alismalllSpotsmenewemcne sje meee tenement L. (N.) amistadensis
16’. Vertex with V-shaped dark mark behind antennal bases; mesoscutum usually with thin transverse dark line along

anterior margin; mesoscutellum completely pale; scape dorsally with two stripes; pronotum with pair of tiny me-

dialispotsxottentiinked tothimidarkepostenorm line ema aa nents acne ne L. (N.) lancala
17(12). Thin dark line extends completely along anterior and lateral margins of pronotum .......... L. (N.) oenops
ie Dark markings on lateral margin of pronotum not extending to anterior margin. ................. 18

18(17'). Pronotum with two medial dark spots; mesonotum with two medial and two anterolateral dark spots
ee se a eae Re ee ee Eo cle Che oo L. (N.) forcipata

18’. Dark spots and stripes on pronotum confined to lateral margins; mesonotal markings restricted to lateral, anterior
aréas ior alliofimesonotims . 21: wes ee DS ee he eg Dee 19
LOGS) Mkronsrandyeenalcomnp le te ly gry alle eames emesis ne ra areca L. (N.) compar
Ih Frons with brownlorred!stripes or spots a. 3 2 = s.sc06 5 4 26) ike 6 oe 6 Ce 20
20(19'). Basal 12 antennomeres very dark; pronotum with thin, very dark brown, lateral stripe. ....... L. (N.) navasi
20'. Basal 12 antennomeres pale yellow, or at most dorsal surface of first six antennomeres darkened; pronotal
markings reddishibrown}thick....(5 fo... Bh elo oe Ee es er Zl

21(20'). Frons with transverse dark stripe below antennal bases; basal six antennomeres dark dorsally; metascutellum dark
L. (N.) nigrovaria

Di Frons with dark spots laterally near gena; antennomeres completely pale; metascutellum pale or dark... .. . 22
22(21’). Pronotum with four reddish brown spots, two laterally oneachside ................. L. (N.) laertes
IPP)! Pronotum with one small reddish brown spot on each anterolateral margin. ................... 23
23i(22)) sMaletsonocomua present, long andithints en. ene) eee eee) eens Seen on ee eee L. (N.) cortesi
DBE Male, SOnOCOMMUA ASEM the eters eitcycen Ranch Suredhcd Gy hve sasGuctemchy bunceeraye Ses pn shes rsetch te) goa RC L. (N.) ratcliffei

Leucochrysa (Nodita) amazonica Navas, 1913
(Figs. 63-65)

DIAGNOSIS. — This is one of four large species of Nodita in Costa Rica with Rs darkened on the
forewing at mid-length. There is a distinctive, red, pentagonal mark on the vertex and an arc of four
red spots along the anterior margin of the mesoprescutum and mesoscutum.

GEOGRAPHICAL DISTRIBUTION. — This species was originally described from Brazil (Amapa,
near the mouth of the Amazon River). Adams (1987) has recorded specimens from El Salvador, Hon-
duras, Panama, and Guyana. Costa Rica records are from: ALAJUELA, Finca San Gabriel, 2 km
southwest of Dos Rios; LIMON, Pandora; Hitoy-Cerere Biological Reserve.

ALTITUDINAL DISTRIBUTION. — Records from 40 to 600 m.

ADULT FLIGHT PERIOD. — March, May, and September.

HABITAT PREFERENCE. — The Pandora and Hitoy-Cerere specimens were collected along small
streams in mature hillside forest on understory saplings.

PENNY: LACEWINGS OF COSTA RICA 197

Leucochrysa (Nodita) amistadensis Penny, 2001
(Figs. 66-68)

DIAGNOSIS. — This medium-sized species of Nodita has dark palpi with their apices pale; an
elongate, dark genal stripe; pale apex to hind wing; a lateral red stripe on both sides of pronotum which
expands beyond mid-length; dark spots at lateral margin of meso- and metascutella; and a darkened
apex of male ninth tergite.

GEOGRAPHICAL DISTRIBUTION. — This species 1s only known from Costa Rica: PUNTARE-
NAS, La Amistad National Park, Las Alturas.

ALTITUDINAL DISTRIBUTION. — 1500-1600 m.

ADULT FLIGHT PERIOD. — February.

HABITAT PREFERENCE. — Collected on understory saplings, including many Peperonia plants,
in mixed montane primary forest with many large Ficus trees.

Leucochrysa (Nodita) antica Navas, 1913
(Figs. 69-71)

DIAGNOSIS. — This is one of three closely-related, medium-sized species of Nodita from Costa
Rica in which the apex of the hind wing is darkened and a lateral red spot is present on both sides of the
pronotum. All three have a curved red line around the anterior margin of the mesonotum and darkened
forewing crossveins, often with margining, especially at the rhegma. All three species have the dorsal
surface of the antennal scape reddened. This species can be separated from the other two by having
two red spots on the frons directly beneath the antennae and having the dorsal red marks wrap around
and cover the ventral surface of the scape.

GEOGRAPHICAL DISTRIBUTION. — The original description of this species indicated only Cen-
tral America for its distribution. Within Costa Rica it has only been collected at: PUNTARENAS, La
Amistad National Park, Las Alturas; Rio Coton.

ALTITUDINAL DISTRIBUTION. — 1500 m elevation.

ADULT FLIGHT PERIOD. — Late February to early March.

HABITAT PREFERENCE. — Collected on understory saplings, including many Peperonia plants,
in mixed montane primary forest with many large Ficus trees.

Leucochrysa (Nodita) askanes (Banks, 1946)
(Figs. 72—74)

DIAGNOSIS. — This is one of two Costa Rican species of L. (Nodita) of medium-size that have
the tips of the hind wings darkened and the posterior half of the pronotum bearing a red triangular
mark laterally on both sides [the other species is Lewcochrysa (Nodita) postica]. Other diagnostic
characteristics include dark maxillary palpi, a transverse red stripe on the clypeus and crossveins that
are not margined in the medial area of the forewing. The vertex bears a small red central spot.

GEOGRAPHICAL DISTRIBUTION. — This species has only been recorded from Costa Rica. Known
localities are: CARTAGO, Turrialba; LIMON, 11.3 km (7 mi) north of Guacimo; Pandora; and
PUNTARENAS, Monteverde.

ALTITUDINAL DISTRIBUTION. — Elevation ca. 40 m at Pandora to 1500 m at Monteverde.

ADULT FLIGHT PERIOD. — February, May, and September with the majority in September.

HABITAT PREFERENCE. — The Pandora locality is a mature wooded hillside with a small stream
running through it. The specimens were collected on the foliage of young understory saplings near the
stream. The Monteverde locality was a more open forest at the edge of a field with individuals again
being found on foliage of understory saplings.

198 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES
Volume 53, No. 12

Leucochrysa (Nodita) caucella (Banks, 1910)
(Figs. 75—77)

DIAGNOSIS. — This medium-sized species has a pale apex to the hind wing; a reddish brown tri-
angular spot laterally on both sides on posterior half of the pronotum and a small reddish brown spot at
its anterio-lateral margin; palpi are dark basally and pale at apex; small dark spot on gena; antennae
pale with dorsal red stripe on scape; mesonotum with two red spots at junction of mesoprescutum and
mesoscutum; forewing radials, gradates, and apical forks dark and with dark margining of gradate
veins.

GEOGRAPHICAL DISTRIBUTION. — This species was previously known from Panama, Colombia,
and Venezuela. Within Costa Rica it is known only from PUNTARENAS, La Amistad National Park,
Las Alturas. This may be a northern South American species that reaches only into southernmost
Costa Rica.

ALTITUDINAL DISTRIBUTION. — 1500—1600 m.

ADULT FLIGHT PERIOD. — February.

HABITAT PREFERENCE. — Collected on understory saplings, including many Peperonia plants,
in mixed montane primary forest with many large Ficus trees.

Leucochrysa (Nodita) compar (Navas, 1921)
(Figs. 78-80)

DIAGNOSIS. — This species has three distinctive characteristics: 1) most characteristic is the dark
brown, basal, ten flagellar segments (apex of flagellum pale); 2) a dark brown, lateral scape stripe, and
3) a thin, dark brown, lateral pronotal stripe. These character states, together with completely pale
frons, gena, and palps, make this species unique among Central American Leucochrysa (Nodita).

GEOGRAPHICAL DISTRIBUTION. — This species was described from Cuba and until now was
known only from that country. Costa Rican specimens (all females) were collected at: ALAJUELA,
20 km south of Upala.

ALTITUDINAL DISTRIBUTION. — Unknown.

ADULT FLIGHT PERIOD. — Late December to early March.

HABITAT PREFERENCE. — This is a lowland species found in disturbed forest habitat.

REMARKS. — The normally dark spot on the forewing pterostigma is quite pale in this species,
the antennae are only slightly longer than the forewing and the outer gradate series meets Psm at a
rather wide angle, leading one to suspect that this may not be a species of Leucochrysini, but rather
Chrysopini (Ceraeochrysa or Chrysopodes). Of the five specimens seen, all were females, so that
male genitalia remain unstudied. Thus, this species is placed here only provisionally.

Leucochrysa (Nodita) cornesta Banks, 1944
(Figs. 81-83)

DIAGNOSIS. — This is one of three, closely-related, medium-sized species of L. (Nodita) from
Costa Rica with the apex of the hind wing darkened and a lateral red spot on both sides of the
pronotum. All three species have a curved red line around the anterior margin of the mesonotum and
darkened forewing crossveins, often with margining, especially at the rhegma, as well as the dorsal
surface of the antennal scape reddened. Leucochrysa (Nodita) cornesta can be separated from the
other two by possession of a complete transverse red line on the frons directly beneath the antennae.
The ventral surface of the antennal scape is pale green.

GEOGRAPHICAL DISTRIBUTION. — This species has previously been known from Surinam and
French Guiana. Within Costa Rica it has been taken at: LIMON, Pandora; Hitoy-Cerere Biological

PENNY: LACEWINGS OF COSTA RICA 199

Station; and PUNTARENAS, Isla. With only four records it is difficult to generalize, but this species
may be limited to the southeastern part of the country.

ALTITUDINAL DISTRIBUTION. — Below 100 m elevation.

ADULT FLIGHT PERIOD. — March and September.

HABITAT PREFERENCE. — This species was collected resting on leaves of understory saplings in
mature hillside forest along a small stream.

Leucochrysa (Nodita) cortesi Navas, 1913

DIAGNOSIS. — This is part of a closely-related group of species with pale maxillary palpi, pale
apex of the hind wing, dark frontal markings, dark lateral markings on the pronotum, and meso- and
metanota that gradually darken with age. Costa Rican species in this group include: L. (N.) cortesi, L.
(N.) laertes, L. (N.) navasi, L. (N.) nigrovaria, and L. (N.) ratcliffei. The most closely-related species
is L. (N.) ratcliffei, which shares with L. (N.) cortesi two dark, lateral, frontal spots rather than a trans-
verse band, a dark spot at the anterolateral corner of the pronotum rather than a lateral stripe, and com-
pletely pale flagellum rather than having the base of the flagellum darkened. Leucochrysa (N.) cortesi
can only be separated from L. (N.) ratcliffei by study of the male genitalia: L. (N.) cortesi has long, thin
gonocornua that are completely absent in L. (N.) ratcliffei.

GEOGRAPHICAL DISTRIBUTION. — This species is recorded from Costa Rica, Mexico (Tabasco),
Trinidad and Venezuela. Within Costa Rica it has been recorded from: LIMON: Hamburg Farm on
the Reventazon (River), (Hamburg Museum—lost); between Limon and Las Mercedes, 12-30 km
from the Atlantic Ocean (Hamburg Museum—lost) [as L. (N.) nevermanni]; SAN JOSE: Pedrogoso,
D.C. Rounds, 1? (MCZ) [as L. (N.) calverti].

ALTITUDINAL DISTRIBUTION. — Sea level (near Limon) to 646 m (at Pedregoso).

ADULT FLIGHT PERIOD. — February, April, and June.

HABITAT PREFERENCE. — Unknown.

REMARKS. — Leucochrysa (N.) calverti and L. (N.) nevermanni, both described from Costa
Rica, were synonymized with L. (N.) cortesi by Banks (1945).

Leucochrysa (Nodita) duarte Banks, 1946 [new combination]
(Figs. 84-86)

DIAGNOSIS. — This is one of three, medium-sized species of L. (Nodita) in Costa Rica witha pale
apex to the hind wing, and red spots along the lateral margin of the pronotum. It can be separated from
the other two by possession of pronotal spots on the posterior half of the pronotum. There is some-
times apparent a second pair of pronotal spots just under the anterio-lateral margin of the pronotum.

GEOGRAPHICAL DISTRIBUTION. — This species is only known from Costa Rica. Records in-
clude: LIMON, Pandora; Hitoy-Cerere Biological Station; and SAN JOSE, Pedregoso (holotype).

ALTITUDINAL DISTRIBUTION. — The Limon Province records are from an elevation of
15-220 m, while Pedregoso is in the central valley at mid-elevations (approximately 1000 m).

ADULT FLIGHT PERIOD. — February and March.

HABITAT PREFERENCE. — This species was collected resting on leaves of understory saplings in
mature hillside forest along a small stream.

Leucochrysa (Nodita) egregia Navas, 1913
(Figs. 87-89)

DIAGNOSIS. — The large size, distinctive red spotting pattern on thorax and abdomen, double red
stripe on vertex, and darkened focal area at mid-point of radial area all help to distinguish this species.

200 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES
Volume 53, No. 12

GEOGRAPHICAL DISTRIBUTION. — This species is known from Guatemala, Costa Rica, and Pan-
ama. Within Costa Rica specimens have been collected at: GUANACASTE, Finca La Pacifica, near
Cafias; and PUNTARENAS, Monteverde.

ALTITUDINAL DISTRIBUTION. — Near sea level to 1500 m.

ADULT FLIGHT PERIOD. — February and September.

HABITAT PREFERENCE. — Specimens were collected at a light trap placed in front of the visitor
center at Monteverde Biological Reserve in mature, moist forest. Individuals have also been collected
in bushes near the river at La Pacifica.

REMARKS. — Leucochrysa (Nodita) luctuosa Banks, 1914 is a new synonym of this species.

Leucochrysa (Nodita) forcipata Penny, 1998
(Figs. 90-92)

DIAGNOSIS. — This species has pale antennae, completely pale frons and gena, two dark spots on
the pronotum and two additional dark spots on the mesonotum. The meso- and metascutella are both
dark. There is a distinctive dark swelling of the first anal crossvein. However, the most characteristic
feature of this species, and the one that gives it its name, is the double dorsal horns of the male
gonarcus.

GEOGRAPHICAL DISTRIBUTION. — Currently, this species is known only from Costa Rica: SAN
JOSE, El Salvage, Rio Tabarcia, 8 km (road) E Palmichal (9.847°N, 84.164°W), 19-21 January 1992,
Holzenthal, Kjer, and Quesada (UMIC).

ALTITUDINAL DISTRIBUTION. — 1650 m.

ADULT FLIGHT PERIOD. — Known only from January.

HABITAT PREFERENCE. — Unknown.

Leucochrysa (Nodita) indiga (Navas, 1928)
(Figs. 93-95)

DIAGNOSIS. — The most distinctive features of this species are: the dark tip of the hind wing, the
dark transverse stripe across the lower part of the frons, the black palpi, the longitudinal dorso-lateral
stripe on the scape, and the darkened first two antennal flagellomeres.

Leucochrysa (Nodita) askanes, L. (N.) indiga, and L. (N.) postica form a closely-related spe-
cies-group with few external characters to separate them. However, L. (N.) postica does not have a
small dark medial spot on the vertex and has extensive dark margining of the forewing inner gradate
crossveins.

Leucochrysa (Nodita) askanes and L. (N.) indiga can only be told apart by characters of the male
genitalia. Leucochrysa (Nodita) indiga lacks the elongate, apically-pointed endoprocesses of the
gonarcus found in both L. (N.) askanes and L. (N.) postica.

GEOGRAPHICAL DISTRIBUTION. — Navas’ original type was indicated as being from Costa Rica:
SAN JOSE, San José. Penny (2001) designated a neotype from ALAJUELA, 20 km south of Upala.
One additional locality is known from LIMON, Pandora.

ALTITUDINAL DISTRIBUTION. — Near sea level.

ADULT FLIGHT PERIOD. — Late September to February.

HABITAT PREFERENCE. — The specimen from Pandora was collected on understory saplings
near a small stream in primary forest where the humidity was quite high and probably remained so the
year round.

PENNY: LACEWINGS OF COSTA RICA 201

Leucochrysa (Nodita) laertes (Banks, 1946)
(Figs. 96-98)

DIAGNOSIS. — This species has two red spots on the frons, often with a thin, inverted, V-shaped
mark connecting them to the area between the antennal bases. There is a thin V-shaped mark on the
vertex behind the antennal bases. A dorso-lateral dark stripe extends the length of the antennal scape
and can, at times, apically cover the whole dorsal surface. The pronotum has a pair of reddish brown
spots laterally on both sides, and pair of tiny brown spots medially. The mesonotum 1s dark laterally
and this can extend along the anterior margin, although it is normally interrupted medially.

GEOGRAPHICAL DISTRIBUTION. — This species has been known only from Panama. Within
Costa Rica this species has been collected only at ALAJUELA, 20 km south of Upala.

ALTITUDINAL DISTRIBUTION. — Lowlands near sea level.

ADULT FLIGHT PERIOD. — Primarily in January, with one specimen collected in May.

HABITAT PREFERENCE. — The only known collecting site from Costa Rica is heavily disturbed,
lowland forest.

Leucochrysa (Nodita) lafoni Navas, 1911

This species was described from an unnamed locality in Costa Rica. The original description
mentions a dark lateral spot on each side of the pronotum and a dark spot on each side of the
mesoscutum. Several species would fit this description, including L. (N.) clypsydra Banks, 1918 from
Colombia; L. (N.) duarte from Costa Rica; and L. (N.) morenoi (Navas, 1934) from Ecuador. The type
(in the Paris Museum) has been eaten out by psocids, is discolored, and lacking an abdomen. At this
time it is probably best to consider this species an incertae sedis.

Leucochrysa (Nodita) lancala Banks, 1944
(Figs. 99-101)

DIAGNOSIS. — The most distinctive characteristics are: the double dark dorsal stripe on the
scape; palpi dark externally; dark red spots on the frons adjacent to gena; dark red V-shaped mark pos-
terior to antennal bases; dark red lateral pronotal stripe; and pair of tiny dark spots medially on the
pronotum. Leucochrysa (Nodita) laertes has many of these features, such as dark spots on the frons,
V-shaped dark mark posteriad to antennal bases; dark scape stripe; and tiny medial pair of spots on the
pronotum, and the two species are probably closely related. However, they differ in that L. (N.) /aertes
has pale palpi, the dark marks on the frons are higher—closer to the antennal bases; the dark scape
stripe is single and thick, often broader apically; and the hind wing apex is pale.

GEOGRAPHICAL DISTRIBUTION. — This species is known from Panama, Colombia, Venezuela
and Surinam. Within Costa Rica it has been collected at: GUANACASTE, La Taboga Forest Reserve,
9 km south of Canas, and La Pacifica, near Cafas.

ALTITUDINAL DISTRIBUTION. — Near sea level.

ADULT FLIGHT PERIOD. — Early to mid-February.

HABITAT PREFERENCE. — Specimens are from lowland, highly modified, riverine forest.

Leucochrysa (Nodita) lenora (Banks, 1944)
(Figs. 102—104)

202 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES
Volume 53, No. 12

forewing crossveins, often with margining, especially at the rhegma. All three species also have the
dorsal surface of the antennal scape reddened. This species can be separated from the other two by its
having two red spots on the frons directly beneath the antennae and lacking red markings on the ven-
tral surface of the antennal scape.

GEOGRAPHICAL DISTRIBUTION. — This species was originally described from Venezuela, Suri-
nam, and Brazil. Costa Rican records include: ALAJUELA, 20 km south of Upala; HEREDIA,
Chilamate; and LIMON, Pandora. This appears to be primarily a northern South American species,
which ranges into southern Central America.

ALTITUDINAL DISTRIBUTION. — The Pandora record is from a hillside only about 40 m above
sea level.

ADULT FLIGHT PERIOD. — All months of the year.

HABITAT PREFERENCE. — This species was collected resting on leaves of understory saplings in
mature hillside forest along a small stream.

Leucochrysa (Nodita) nativa (Navas, 1911)

This Costa Rican species is rather large, with a forewing length of 17.5—18 mm. It is clearly a
member of Leucochrysa (Nodita), but the original description is not very informative and the type is
badly damaged. The specimen (MNHN, Paris) is badly discolored, the head is crushed, and it is cov-
ered with moth scales. It is best to consider it a nomen dubium at this time.

Leucochrysa (Nodita) navasi Kimmins, 1940
(Figs. 105-107)

DIAGNOSIS. — This is one of the medium-sized species of L. (Nodita) with pale palpi and pale
apex to hind wing; a lateral dark stripe on pronotum which ends at anterlateral angle. This species can
be separated from other species in this group by the small, reddish brown spot on frons below antenna;
completely pale vertex; meso- and metanota completely pale; forewing veins darkened at mid-length
of radius, on gradates and along apical forks.

GEOGRAPHICAL DISTRIBUTION. — This species was originally described from Mexico under the
junior homonym Nodita alternata. There is an unpublished record from Honduras. Within Costa Rica
it has been collected at: CARTAGO, Cachi; PUNTARENAS, La Amistad National Park, Las Alturas;
Fila Cedra, near Las Alturas.

ALTITUDINAL DISTRIBUTION. — 1500-2000 m.

ADULT FLIGHT PERIOD. — February.

HABITAT PREFERENCE. — Collected on understory saplings, including many Peperonia plants,
in mixed montane primary forest with many large Ficus trees.

Leucochrysa (Nodita) nigrovaria (Walker, 1853)
(Figs. 108—110)

DIAGNOSIS. — This medium-sized species has the apex of the hind wing pale; pale maxillary and
labial palpi, a transverse dark stripe on the frons below the antennae; vertex bearing a red V-shaped
mark behind the antennae; pale pronotum with dark lateral margin and variably darkened meso- and
metanota. The antennae are dark basally and progressively paler after the basal six flagellomeres.

GEOGRAPHICAL DISTRIBUTION. — This species has been recorded from Colombia, Venezuela
and Costa Rica (unnamed locality) under the name L. (N.) ceratica. The only other record is from:
PUNTARENAS, La Amistad National Park, Las Alturas.

PENNY: LACEWINGS OF COSTA RICA 203

ALTITUDINAL DISTRIBUTION. — The Las Alturas record is from 1500 m elevation, and the area
near San José is at mid-elevations (1000-1200 m).

ADULT FLIGHT PERIOD. — March.

HABITAT PREFERENCE. — Collected on understory saplings, including many Peperonia plants,
in mixed montane primary forest with many large Ficus trees.

REMARKS. — This species has slowly developing, dark pigmentation on the meso- and
metanota. The first dark pigment appears on the metascutellum, then progressively develops on the
rest of the meso- and metanota. Leucochrysa (N.) ceratica (Navas, 1911) and L. (N.) maculata (Navas,
1928) are new synonyms.

Leucochrysa (Nodita) oenops Adams, 1987
(Figs. 111-113)

DIAGNOSIS. — The most distinctive feature is a narrow dark ring around the lateral and anterior
margins of the pronotum. Many other species have lateral stripes, but none extend fully around the an-
terior margin, as does this species. This is one of the medium-sized species of Nodita in Costa Rica
with pale palpi and pale apex to the hind wing. Two other features help set this species apart from oth-
ers: behind the antennae on the vertex are double V-shaped red marks followed by a triangular red cen-
tral mark on the vertex. The mesonotum is variable, having at minimum a pair of anterior dark spots
and maximally having the whole mesonotum darkened.

GEOGRAPHICAL DISTRIBUTION. — At present this species is only known from Nicaragua and
Costa Rica. Costa Rican records include: GUANACASTE, La Taboga Forest Reserve, 9 km south of
Cafias; 14 km south of Cafias; Finca La Pacifica, near Canas; Santa Rosa National Park; and LIMON,
Pandora.

ALTITUDINAL DISTRIBUTION. — All known localities are lowland, with the Pandora locality at
about 40 m.

ADULT FLIGHT PERIOD. — March, May, and September.

HABITAT PREFERENCE. — Individuals of this species were collected resting on leaves of
understory saplings in mature hillside forest along a small stream at Pandora. Other individuals were
collected in bushes near a river at La Pacifica.

Leucochrysa (Nodita) pallescens Banks, 1946
(Figs. 114-116)

DIAGNOSIS. — This is one of the four large species of L. (Nodita) in Costa Rica with a dark focal
area in the radial area of the forewing. There is a thin, dark, V-shaped marking on the vertex behind the
antennae and only occasional, very small markings on the thorax; an occasional small, dark
right-angled mark at the posterior part of the pronotum and a thin dark line tranversely across the ante-
rior mesoscutum.

GEOGRAPHICAL DISTRIBUTION. — This species is only known from Guatemala and Costa Rica.
Within Costa Rica it is known only from: LIMON, Pandora.

ALTITUDINAL DISTRIBUTION. — Pandora is only about 40 m above sea level.

ADULT FLIGHT PERIOD. — September.

HABITAT PREFERENCE. — The Pandora specimen was collected on understory saplings in ma-
ture hillside forest along a small stream.

204 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES
Volume 53, No. 12

Leucochrysa (Nodita) postica Navas, 1913
(Figs. 117-119)

DIAGNOSIS. — This is one of two medium-sized species of L. (Nodita) with a darkened hind wing
tip and a triangular red mark laterally on both sides of the posterior half of the pronotum [the other is
Leucochrysa (Nodita) indiga]. The maxillary palpi are dark; the clypeus bears a transverse red stripe;
and the m-cu crossveins are margined in the forewing. The vertex is completely pale green.

GEOGRAPHICAL DISTRIBUTION. — This species has previously been reported from Surinam and
French Guiana. Within Costa Rica, records are from: ALAJUELA, 20 km south of Upala;
GUANACASTE, 20 km southwest of Canas; HEREDIA, La Selva Biological Station, near Puerto
Viejo; LIMON, Hitoy-Cerere Biological Reserve; and PUNTARENAS, 16.1 km(10 mi) S Rincon de
Osa.

ALTITUDINAL DISTRIBUTION. — Below 200 m.

ADULT FLIGHT PERIOD. — Most records (n= 10) are from February to May, but there is one re-
cord from September.

HABITAT PREFERENCE. — The collections made at Hitoy-Cerere Biological Reserve were on
understory saplings in mature hillside forest along a small stream.

Leucochrysa (Nodita) ratcliffei Penny, 2001
(Figs. 120-122)

DIAGNOSIS. — This is amedium-sized species of L. (Nodita) with pale hind wing apex; pale pal-
pi; small brown spot on frons between gena and antennal base; dark spot on vertex at dorsal base of an-
tenna; small brown spot at anterio-lateral margin of pronotum; dark arc along anterior margin of
mesonotum; forewing gradates, apical forks, and rhegmal spot darkened and margined.

GEOGRAPHICAL DISTRIBUTION. — Known only from Costa Rica. Within the country it has been
collected at: LIMON, Pandora; and PUNTARENAS, Las Cruces, near San Vito.

ALTITUDINAL DISTRIBUTION. — The Pandora record is from lowland hillside at about 100 m ele-
vation, while the Las Cruces records are from near 1800 m elevation.

ADULT FLIGHT PERIOD. — March and September.

HABITAT PREFERENCE. — Individuals of this species were collected resting on leaves of
understory saplings in mature hillside forest along a small stream.

Leucochrysa (Nodita) serrei (Navas, 1923)
(Figs. 123-125)

DIAGNOSIS. — This medium-sized species’ most notable characteristics are the pale green, un-
marked pronotum and the reddish dorsal surface of the antennal scape. No other species of L. (Nodita)
in Costa Rica has an unmarked pronotum. The male gentialia also have a distinctive pair of dorsal
horns on the arcessus, which are blunt-tipped with a U-shaped incision between them.

GEOGRAPHICAL DISTRIBUTION. — This species has only been collected in Costa Rica. The re-
cords are: GUANACASTE, 14 km south of Cafias; San Gabriel; Maritza Biological Station, west side
of Orosi Vulcano; SAN JOSE, San José.

ALTITUDINAL DISTRIBUTION. — 600 m.

ADULT FLIGHT PERIOD. — March and July.

HABITAT PREFERENCE. — Unknown.

PENNY: LACEWINGS OF COSTA RICA 205

Leucochrysa (Nodita) superior Navas, 1913

DIAGNOSIS: This is a fairly large species with dark maxillary palpi, dark clypeus, dark lateral
margins of pronotum, and two mesonotal spots. The most similar species is L. (N.) caucella, which
can be separated by its smaller size and pale clypeus.

GEOGRAPHICAL DISTRIBUTION. — The type is from Guatemala. Navas (1928) reported this spe-
cies from COSTA RICA: LIMON, between Limon and Las Mercedes, 12—30 km from the Atlantic
Ocean (Hamburg Museum—lost).

ALTITUDINAL DISTRIBUTION. — Sea level.

ADULT FLIGHT PERIOD. — November.

HABITAT PREFERENCE. — Unknown.

REMARKS. — This is a poorly known species, with a single, somewhat dubious record from
Costa Rica. The type is teneral and may not display the full coloration of a mature individual.

Leucochrysa (Nodita) virginiae Penny, 1998
(Figs. 126-128)

DIAGNOSIS. — This medium-sized species has the apex of the hind wing pale; pale maxillary and
labial palpi; a completely red frons; pale vertex; pale pronotum and metanotum.

GEOGRAPHICAL DISTRIBUTION. — This species has only been collected in Costa Rica. The only
record is from: LIMON, Hitoy-Cerere Biological Reserve.

ALTITUDINAL DISTRIBUTION. — 220 m.

ADULT FLIGHT PERIOD. — March.

HABITAT PREFERENCE. — This species was collected resting on leaves of understory saplings in
mature hillside forest along a small stream.

Vieira Navas, 1913

This is one of the most distinctive of chrysopid genera. In addition to being among the largest in
size and having a very characteristic dark pigmentation pattern on the wings, the costal area of the
forewings is very broad with sinuous basal crossveins. This is the only genus with costal crossveins of
this form.

There are currently two recognized species in the genus, but they are possibly synonymous
(Brooks and Barnard 1990). There is an additional undescribed species in northern South America.
The genus ranges from Mato Grosso State in west central Brazil to Costa Rica. One species, Vieira
leschnaulti, has been collected in Costa Rica.

Vieira leschnaulti (Navas, 1911)
(Figs. 129-131)

DIAGNOSIS. — The large size, extensively pigmented forewings, and sinuous basal costal veins
serve to distinguish this species from other Chrysopidae found in Costa Rica.

GEOGRAPHICAL DISTRIBUTION. — This species was originally described from French Guiana,
and has been found from Costa Rica and Surinam south to west central Brazil, Peru and Bolivia.
Within Costa Rica, only three collections are known: CARTAGO, Centro Agronomico Tropical de
Investigacion y Ensenanza (C.A.T.I.E.), 2 km southwest of Turrialba (twice); and LIMON, Cerro
Cocori, Finca de E. Rojas.

ALTITUDINAL DISTRIBUTION. — Unknown.

206 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES
Volume 53, No. 12

ADULT FLIGHT PERIOD. — The collections at C.A.T.I.E. were in late May and late June while the
Cerro Cocori specimen was collected in February.

HABITAT PREFERENCE. — The specific habitat where the Costa Rican specimens were collected
is unknown, but in the central Amazon Region a specimen was collected on understory vegetation ina
mature forest.

NOTE. — It is probably a coincidence that the two largest and most colorful species of New
World chrysopids, Loyola croesus and Vieira leschnaulti, are both seldom collected, yet within Costa
Rica both have been collected in the immediate vicinity of Turrialba during May and June.

TRIBE BELONOPTERYGINI

This group of green lacewings is very robust, with broad bodies, pronota that are wider than long,
and very stout antennae with flagellar segments wider than long. Very little is known of the biology of
this tribe, but where life history has been studied, the larvae are always associated with ant nests. One
blue-colored larva of Nacarina was encountered in the nest of Camponotus ants in Guyana. The Old
World genus /talochrysa is speciose, but in the Western Hemisphere there are only three genera and
16 known species (Brooks and Barnard 1990). The latter range from Cuba and the southeastern
United States to northern Argentina. Only Nacarina has been recorded from Costa Rica.

Nacarina Navas, 1915

Nacarina is easily separated from the other two New World genera of Belonopterygini by the ab-
sence of dark brown or black pigmentation on the body and wings (as in Abachrysa) and having nei-
ther the narrow wings, nor the longitudinal red stripe of the forewing, of Belonopteryx. Nacarina
contains 14 of the 16 known species of New World Belonopterygini. They are found over the same
range as the tribe in the New World, that is, from the southeastern United States to Argentina. Three
species have been collected in Costa Rica.

KEY TO COSTA RICAN NACARINA

1. Three rows of gradate crossveins in forewing; forewing >30mminlength .................. N. titan
i) Iwoxows'of gradate crossveins im forewing: forewing. <30 mmiinilength, | >=... . . sen 2
2(1'). Antennae entirely very dark; body brown and forewing longitudinal veins black ............ N. megaptera
2'. Antennae pale or possibly with exterior dark stripe; body green, with forewing longitudinal veins green (some
crossvermsidarle)s : Path Sit Aas PP ITE NE et Ee ES BOL Ee RE Ee eS | ee N. cordillera

Nacarina cordillera (Banks, 1910)
(Figs. 132-134)

DIAGNOSIS. — This is the most common species of Nacarina in Costa Rica, and perhaps in the
New World. It is frequently attracted to lights and often appears in collections. Only recently was the
synonymy with N. balboana recognized and most identified specimens used the latter name. This spe-
cies has a pale green body and wings with some crossveins darkened, and antennae pale (often with
dark lateral stripe), with only two rows of gradate crossveins. The male ninth sternite has an
apico-medial protuberance.

GEOGRAPHICAL DISTRIBUTION. — This species is known from southern Mexico, El Salvador,
Costa Rica, Panama, Colombia, Venezuela, and western Brazil (Rond6nia). Within Costa Rica this
species appears to be commonly distributed in all lowland areas of the country and is readily attracted
to lights. It has been collected at: ALAJUELA, Fabio Baudrit M. Agricultural Experiment Station;

PENNY: LACEWINGS OF COSTA RICA 207

CARTAGO, 7km east of Moravia; Turrialba; Playa del Coco; 2km southeast of Tuis;
GUANACASTE, Santa Rosa National Park (PNSR), Rio Cuajiniquil; PNSR, Rio Poza Salada;
PNSR, Quebrada Guacimo; PNSR, Quebrada Pedregal; PNSR, Agua Buena; Cerro El Hacha; west
side of Orosi Vulcano, Maritza Biological Station; La Pacifica, 6 km north of Canas; Murciélago Bio-
logical Station, 8 km southwest of Cuajiniquil; Pitilla Biological Station, 9 km south of Santa Cecilia;
Palo Verde Biological Station, Palo Verde National Park; Barra Honda National Park, 3 km northwest
of Nacaome; Las Pailas Biological Station, Rincon de la Vieja National Park; Finca Jenny, 30 km
north of Liberia; HEREDIA, Magsasay Biological Station, Braulio-Carrillo National Park; LIMON,
Finca E. Rojas, Cerro Cocori Sector; Cuatra Esquinas Biological Station, Tortuguero National Park;
Cerro Tortuguero, Tortuguero National Park; Hitoy-Cerere Biological Station, Hitoy-Cerere Biologi-
cal Reserve; Estrella Valley, Hitoy-Cerere Biological Reserve; Manzanillo, Gandoca y Manzanillo
National Wildlife Refuge; Rio Sardinas, Barra del Colorado National Wildlife Refuge;
PUNTARENAS, Quebrada Bonita Biological Station, Carara Biological Reserve; Quepos, Manuel
Antonio National Park; Rancho Quemado, Osa Peninsula; Bosque Esquinas, Osa Peninsula.

ALTITUDINAL DISTRIBUTION. — Sea level to 1050 m, but most common at lower elevations.

ADULT FLIGHT PERIOD. — January to July.

HABITAT PREFERENCE. — Collection records from Costa Rica indicate dry, semiarid environ-
ments, while Brazilian records indicate heavily disturbed pasture lands.

Nacarina megaptera (Navas, 1927)
(Figs. 135-137)

DIAGNOSIS. — This species is recognized by the completely dark antennae, brownish body, dark
wing venation and two rows of gradate veins. The male ectoproct is ventrally pointed.

GEOGRAPHICAL DISTRIBUTION. — Originally described from Brazil, this species is also known
from Costa Rica and Guyana. Within Costa Rica it has been collected at: GUANACASTE, Las Pailas
Biological Station, Rincon de la Vieja National Park; Agua Buena, Guanacaste National Park;
HEREDIA, Magsasay Biological Station, Braulio-Carrillo National Park; LIMON, Tortuguero Na-
tional Park, 3.5 km south of Tortuguero; Rio Sardinas, Barra del Colorado Wildlife Reserve;
Hitoy-Cerere Biological Reserve, PUNTARENAS, Carara Biological Reserve (RBC), Quebrada
Bonita Biological Station; RBC, El Chuzazo; Corcovado National Park, Piedra el Arco; Quepos,
Manuel Antonio National Park; Esquinas Biological Station, Osa Peninsula. It appears to be fre-
quently attracted to lights at Manuel Antonio National Park, especially during April.

ALTITUDINAL DISTRIBUTION. — Sea level to 800 m.

ADULT FLIGHT PERIOD. — Collection records exist for almost all months of the year, but are con-
centrated in two modalities from February to June, and August to December.

HABITAT PREFERENCE. — This species appears to be taken at lights in the same habitat with N.
cordillera.

Nacarina titan Banks, 1915
(Figs. 138-140)

DIAGNOSIS. — This species is characterized by its very large size (more than 30 mm forewing
length), pale coloration, and disorganized third (middle) row of gradate crossveins. The pronotum has
spots rather than stripes. The male ninth sternite bears apically a bifurcate protuberance.

GEOGRAPHICAL DISTRIBUTION. — The type specimen of Nacarina titan was collected, at Costa
Rica: LIMON, Limon. A second specimen is known from Hitoy-Cerere Biological Reserve, south of
the town of Limon.

208 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES
Volume 53, No. 12

ALTITUDINAL DISTRIBUTION. — The port city of Limon is close to sea level. The specimen from
Hitoy-Cerere Reserve was collected at 200 m elevation.

ADULT FLIGHT PERIOD. — Collection dates are late May and December.

HABITAT PREFERENCE. — Unknown.

TRIBE CHRYSOPINI

This is the predominant group of green lacewings in the world, and perhaps the largest tribe of all
neuropterans. They are ubiquitous in temperate regions where they sometimes go by the names of
golden-eyes, or stink bugs, although the term “stink bugs” is more correctly reserved for the
hemipteran family Pentatomidae. They have been reared for mass introductions as biological control
agents in fields and orchards. Chrysopines have been largely replaced in the New World tropical for-
ests by leucochrysines, but they remain dominant in tropical open areas, including most agricultural
lands. There are 32 known genera of Chrysopini worldwide, five of which are found in Costa Rica.

Ceraeochrysa Adams, 1982a

Ceraeochrysa is a New World genus of chrysopine green lacewings containing about 50 species
(see Adams 1982a). Although several species are known from temperate areas, most species are trop!-
cal. Costa Rica has a very rich assemblage of this genus, with 20 known species found within its bor-
ders. They are one of the most commonly encountered groups of green lacewings in the New World
tropics. The habitat is open areas, such as grasslands, orchards, plantations, urban areas and cultivated
plots, and they are seldom found in deep forest. Because of their presence in agricultural habitats,
many species have good potential as biological control agents.

In general habitus they are medium-sized (about 10—15 mm in forewing length) with relatively
narrow wings bearing little or no dark pigmentation on the membrane. However, the head and thorax
are often brightly patterned in red or brown on a pale green background. Most Costa Rican species
have a lateral longitudinal stripe on the pronotum, although some species have lateral spots on the
pronotum. Unfortunately, the subcutananeous pigments break down rather easily and quickly after
death, so that most specimens that have been exposed to light, heat, or chemicals for any length of time
bear little resemblance to the colorful live adult.

The most characteristic feature of this genus is the very elongate gonapsis of the male genitalia.
Except for a few species of Chrysopodes with short, broad gonapses, no other New World chrysopids
have this distinctive form of genital armature.

Larvae of this genus are known to be trash carriers. That is, they place debris and the dried bodies
of their prey over their own bodies, which are modified with long setae to hold the trash.

KEY TO COSTA RICAN CERAEOCHRYSA

ile Lateral surface ofipronotum withispots;/antennaeand palpi pale 2. 5 = 5. = «2 4) .9)-)sen een 2,
IVY. Pronotum withilateralistnipes:)antennaciand)palpivaniable ss S)2)-e asses) ee) See ee 8
2()S) Mesoscutum: wathitwordark:centrallspots) 9) 2 = 0 25) oe 2 5 ee 5
Pais Mesoscutum completely pale") oe ake me ews eee ee se ee se os 60 Gc 3
3(2))3 Forewing venationiwathoutsswollens heayilymarcined arcasyar- ete alate ee ene nee eee o
3% Forewing with swollen and darkly margined 2A-3A crossvein and apex ofCuP ............ C. tauberae
4(3). One large dark spot anterior to mid-length on each side of pronotum ................ C. costaricensis

4". Two large dark spots on each side of pronotum, one of these at posterio-lateral angle ......... C. defreitasi

PENNY: LACEWINGS OF COSTA RICA 209

5(2). Metanotum with a pair of dark spots; apex of forewing CuP swollen, darkened... ............... 6
Sf. Metanotum pale, without dark markings; apex of forewing CuP not swollen ................... aT
6(5). | Antennal scape without a dark stripe; vertex not darkened; apical tarsal segment dark. ........ C. nigripedis
6’. Antennal scape with a dark stripe dorsally; vertex darkened medially; apical tarsal segment pale . . . . C. elegans
7(5'). Two pairs of dark lateral pronotal spots; forewing crossvein 2A-3A not swollen and darkened ..... . C. inbio
he One pair of dark lateral pronotal spots; forewing crossvein 2A-3A swollen and darkened ....... C. discolor

8(1'). Inner gradate series of forewing bearing a small reticulate mosaic of cells surrounded by darkened membrane
ee ee ee Le nets ee ee ee kee et ee oe C. pseudovaricosa

oe Inner gradate series normal, with stepwise series of veins, without reticulate mosaic of small cells ....... 9
Ui ee Bascomantconalbilagellumpaleon dorsalisuntacel aa) ene eeceeiem-s) sation) tcl tN aicl ene sce 10
OV. Base Cr amiga mbyeelivion Chrde@iolopell sbineice S95 5 a 55 45 soo eee he ee ee ee ee 13
10(9). Pronotal stripes narrow, only slightly wider than antennal flagellum; two narrow dorsal stripes on antennal scape .
ere (COntimucmonlywithtmales)M ett ae. ec) etc ee eee ne icles sve irae ais bs Soe eens 11
10’. Pronotal stripes broad, two to three times as wide as antennal flagellum; a single dorsal or lateral stripe on antennal
SOF 2c (COMMS OA Winns) a oo oa eo ee oh Oooo doch aoe oOo hE Oe 12
11(10). Male ninth sternite forming a recurved apical horn juxtaposed against numerous stout denticles ventrad of
POUAD SIS EMR ee i hoon cere eter ef ee ee, choot emirate «heen ema ets C. arioles
Wily Maleminthistemite notxecurved dorsally asjantapical’ hom) 22). 55-2. 4.06565 2) ee oe C. cincta
12(10’). Male abdomen with very enlarged spiracles; dorsolateral lobe absent from ninth sternite, but apex bifurcate and
dentate, with additional spines lateral to apex of gonapsis; dorsal lobe of gonarcus singular, medial . . . C. cubana
ae Male abdomen bearing normal spiracles; dorsolateral lobe present on ninth sternite, apex rounded with paired
elongate ridges on membrane above ninth sternite; dorsal lobe of gonarcus paired, caudally recurved . . C. valida

13(9'). Dorsal surface of antennal scape completely red; vertex at base of antennal scapewith broadly triangular red mark
PE OR ie Ry ties be ies EA al eS He Pals ai Ne santo ey Me CE er ee es C. smithi

13 Dorsal surface of antennal scape pale or with longitudinal stripes, but not completely red; antennal marks may
extend onto vertex as thin stripe, but not as broadly triangularmark ..............--.--..--.. 14

lon) sDarkieenal line present between compound (eye and labrum’ fs) | | = ee IS)

14". Genal area below eye pale or with only a diffuse red tinge. . . . (continue only with males) .......... 16

15(14). Pronotal stripe reddish brown; marginal forks of forewing pale green; pterostigma completely pale green to

yellow; setae on male ectoproct slender, not stalked; gonarcus with medialhood ............. C. everes
Se Pronotal stripe chocolate brown; marginal forks of forewing darkened; pterostigma with slight darkening

anteriorly; setae on male ectoproct thickened, stalked; gonarcus without medial hood ......... C. berlandi
16(14’). Dorsal apodeme of male ectoproct with ventral anvil-shaped expansion .........-......--- C. acmon
16’. Dorsal apodeme of male ectoproct without ventral anvil-shaped expansion... .....-..--..--5-55 17

17(16'). Pronotal stripes much broader than antennal flagellum; male ninth sternite with dorsolateral projection and two
parallel ridges on membrane above apex; male genitalia with heavily sclerotized teeth on gonosaccus and broad

enLoplocesses wilich taper rapidlyzatiapexemane y snei eae cee eer ek Gen = ee ewe eee cee C. sanchezi
rae Pronotal stripes about same width as antennal flagellum; male ninth sternite without dorsolateral projection or par-

allel ridges above apex; male genitalia without stout teeth on gonosaccus. ...........2.-252255,5 18
18(17'). Antennal scape with dorsolateral stripe halfas wide aslong ............--.+++--+-+--. C. caligata
18". Antennal scape with one dorsal and often one lateral stripe one-quarter as wide aslong ....-.....-.-. 19

19(18'). Pronotal stripe brown; outer gradate series of forewing with seven or fewer crossveins; head yellow, distinctly
COMMANIINS Win MSN HOMO » 5 oueacnaons sear eeanosndaoecno deen moo me C. claveri
IS Pronotal stripe red; outer gradate series of forewing with nine or more crossveins; head and pronotum concolorous
ee re, ey oh OR le eas an rey IE opiate see 58, Syee: SO mune ee eee ae C. gradata

210 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES
Volume 53, No. 12

Ceraeochrysa acmon Penny, 1998
(Figs. 141-146)

DIAGNOSIS. — This species is similar to several other Costa Rican species of Ceraeochrysa with
dark antennae and lateral red stripe on the pronotum. The most distinctive feature is the anvil-shaped
ventral extension of the dorsal apodeme of the male terminalia. Identification without resort to study
of the male terminalia is not reliable.

GEOGRAPHICAL DISTRIBUTION. — Ceraeochrysa acmon has, until now, only been recorded
from Costa Rica, Panama, and Brazil. Within Costa Rica, this species is known from the holotype at:
LIMON: Amubri, A. C. Amistad, UTM coordinates L-S 385500, 578000, 70 m, 4-21 December
1993, G. Gallardo, | male (INBIO), and GUANACASTE, Santa Rosa National Park.

ALTITUDINAL DISTRIBUTION. — This appears to be a lowland species. The record from Amubri
was only 70 m and Santa Rosa National Park is near sea level. The site near Ariquemes in Brazil is at
about 200 m elevation.

ADULT FLIGHT PERIOD. — In Costa Rica specimens were collected in April and December, but
specimens from Panama were collected in May, and Brazilian specimens in October and November.

HABITAT PREFERENCE. — The site where this species was collected near Ariquemes in Brazil is
secondary forest and abandoned plantations. There is also pasture land nearby. Santa Rosa National
Park is primarily dry, deciduous tropical forest.

Ceraeochrysa arioles (Banks, 1944)
(Figs. 147-152)

DIAGNOSIS. — This species has pale labial palpi and antennae; two dorsal red stripes on antennal
scape; pronotal stripes red and narrow; forewing gradates dark—3 inner gradates and 5 outer gradates.
The most characteristic feature is the male terminalia, which includes a unique, recurved horn at the
apex of the ninth sternite.

GEOGRAPHICAL DISTRIBUTION. — This species was originally described from San José, Costa
Rica (Navas 1928). In the literature this species has been recorded from Mexico (Chiapas, Nuevo
Leon, and Sinaloa), Honduras, Guatemala, and Trinidad under the names C. binaria and C. arioles.
There is a female specimen determined by Nathan Banks from Guyana in the Field Museum (Chi-
cago), although females are currently inseparable from C. cincta and C. caligata, and earlier records
based on females must be viewed with much caution. At the time Adams (1982a) erected the genus
Ceraeochrysa he designated a neotype for this species from Chiapas, Mexico. There is, in the Utah
State University and California Academy of Sciences, a series of specimens collected at SAN JOSE:
Escazu, a suburb of Costa Rica’s capital city.

ALTITUDINAL DISTRIBUTION. — The elevation of Escazu is 1100 m.

ADULT FLIGHT PERIOD. — The Escazu specimens were collected during the months of January,
April, May, and July.

HABITAT PREFERENCE. — Collections from the urban Escazut site indicate an open, disturbed
habitat preference.

REMARKS. — This species was originally described as Chrysopa binaria Navas 1928 (a junior
homonym of Chrysopa binaria Navas 1923) from San José, Costa Rica. The type was deposited in the
Hamburg Museum and subsequently destroyed during WWII. Banks (1944) recognized the
homonymy and created the name Chrysopa arioles.

PENNY: LACEWINGS OF COSTA RICA PAI

Ceraeochrysa berlandi (Navas, 1924)
(Figs. 153-158)

DIAGNOSIS. — This is a member of the /ineaticornis group of species with dark antennae, dark
brown lateral scape stripe, a thin dark lateral stripe on the pronotum, and male terminalia containing
an elongate, apically tripartite mediuncus, apically tapered ectoproct, gonapsis that is apically bifur-
cate, and a pair of erect, elongate gonocornu at the base of the mediuncus.

This species can be separated from other members of this group by the thickened setal bases at the
apex of the ectoproct and the extremely thickened setal bases apically on the ninth sternite. This is one
of two known species of Ceraeochrysa in Costa Rica with heavily margined forewing crossveins (the
other being C. gradata). Additional characters include a dark brown genal spot, pale palpi, a dark
brown lateral stripe on the antennal scape, and the first 10 to 12 antennal flagellomeres dark; apical
antennal flagellomeres pale. The most distinctive features are the sclerites of the male genital arma-
ture, including: chalazate setae at the apex of the ectoprocts and ninth sternum; large, strongly curved
entoprocesses; and two large lateral patches of gonosetae.

GEOGRAPHICAL DISTRIBUTION. — This species was described from Costa Rica, without specific
locality information. There is a specimen in the Museum of Comparative Zoology, Harvard Univer-
sity from Mexico (Chiapas). Within Costa Rica specimens are known from CARTAGO, Tapanti Na-
tional Wildlife Refuge; and PUNTARENAS, Osa Peninsula, Rancho Quemado.

ALTITUDINAL DISTRIBUTION. — Mid-elevation at Tapanti to low elevation at Rancho Quemado.

ADULT FLIGHT PERIOD. — Both records from Costa Rica are from January.

HABITAT PREFERENCE. — Unknown.

Ceraeochrysa caligata (Banks, 1946)
(Figs. 159-164)

DIAGNOSIS. — This is one of the species of Ceraeochrysa with pale antennae. It has a relatively
narrow pronotal stripe and narrow stripe dorsally on scape. This species 1s most closely related to C.
cincta and can be separated from it most easily by studying the male genitalia which have longer lat-
eral arms of the gonarcus and the gonocornu are much longer than the mediuncus horns.

GEOGRAPHICAL DISTRIBUTION. — This species was originally described from Panama. Adams
and Penny (1987) reported this species from the Amazon Basin of Brazil. Within Costa Rica C.
caligata is known from GUANACASTE: 8 km (5 mi) NE Canias; Finca La Pacifica..

ALTITUDINAL DISTRIBUTION. — Lowland areas below 200 m.

ADULT FLIGHT PERIOD. — Late December and early February.

HABITAT PREFERENCE: Two males were collected in bushes near a river in early February.

Ceraeochrysa cincta (Schneider, 1851)
(Figs. 165-170)

DIAGNOSIS. — As with C. caligata and C. arioles, this species has pale labial palpi, pale anten-
nae, red stripe dorsally on antennal scape; and narrow red pronotal stripe. The male genitalia bear a
rather curved ninth tergite apodeme, short entoprocesses, and short gonocornu.

GEOGRAPHICAL DISTRIBUTION. — This species was originally described from Brazil, and Ad-
ams and Penny (1987) recorded it from the southern United States, Mexico, Cuba, Guatemala, Pan-
ama, Galapagos Islands, Guyana, Surinam, Peru, Uruguay, and Argentina. It has also been collected
frequently in northern Costa Rica from: ALAJUELA: 20 km S Upala; and GUANACASTE: 14 km S
Canas, 8 km (5 mi) NE Camas, 3 km SE Ri6 Naranjo.

ALTITUDINAL DISTRIBUTION. — In Costa Rica known only from below 200 m.

PENNY: LACEWINGS OF COSTA RICA 12

ADULT FLIGHT PERIOD. — A possible emergence peak from October to April. This is the driest
time of year for most of Costa Rica.

HABITAT PREFERENCE. — Known records in Costa Rica are from the lowland farming areas and
disturbed secondary forest. Near Canas, in February, several adults of this species were found resting
on bushes near a river.

Ceraeochrysa claveri (Navas, 1911)
(igs 171176)

DIAGNOSIS. — This species has pale palpi and very dark antennae. There is a double stripe on the
antennal scape but it is placed dorsolaterally, so that the most lateral stripe is on the lateral surface. The
pronotal stripe is narrow and brown. There are three to four inner and six to seven outer gradate veins
in the forewing. The male terminalia have a curved hook on the ectoproct apodeme and there is a dis-
tinctive bifurcated apex to the male gonapsis.

GEOGRAPHICAL DISTRIBUTION. — Adams and Penny (1987) record this species from Mexico,
Cuba, Haiti, Puerto Rico, Guatemala, Honduras, Panama, Colombia, Trinidad, Guyana, Surinam and
Brazil. Within Costa Rica this species has been collected at: CARTAGO, Rancho Naturalista, NE
Tuis; GUANACASTE, 8 km (5 mi) NE Canfas; HEREDIA, Puerto Viejo, Rio Sarapiqui, Finca La
Selva; and LIMON, Sector Cerro Cocori, Finca de E. Rojas; and Pandora.

ALTITUDINAL DISTRIBUTION. — 100 to 150 m in the lowlands, and somewhat higher elevations
at Rancho Naturalista.

ADULT FLIGHT PERIOD. — February, June, September, and December in Costa Rica.

HABITAT PREFERENCE. — Ceraeochrysa claveri was attracted to lights near residences in an
open area with many ornamental plants and virtually surrounded by banana plantations.

Ceraeochrysa costaricensis Penny, 1997
(Figs. 177-182)

DIAGNOSIS. — This species has pale green to yellow features, except for a pair of dark spots on
the pronotum. In this respect it differs from other species of Ceraeochrysa with a spotted pronotum, in
that they have four spots on the pronotum and dark spots on the mesoscutum and sometimes
metascutum. In addition, there is no swelling and darkening of veins at the apex of the posterior
cubitus of the forewing. The male genitalia indicate a close relationship with C. everes, which has a
longitudinal red stripe laterally on both sides of the pronotum.

GEOGRAPHICAL DISTRIBUTION. — This species is known from the type male collected at Costa
Rica: PUNTARENAS, Los Alturas, 27 February 1991, Helen Sparrow (INBIO); and Las Cruces,
6 km S San Vito.

ALTITUDINAL DISTRIBUTION: 1360 m.

ADULT FLIGHT PERIOD. — Late February and late September to early October.

HABITAT PREFERENCE. — The holotype specimen was attracted to a kerosene lamp inside a
building at Los Alturas. The surrounding area is almost entirely disturbed, cultivated land, including
ornamental trees around homes and orchards. The Las Cruces locality is the Wilson Botanical Garden
only a few kilometers from the type locality.

Ceraeochrysa cubana (Hagen, 1861)
(Figs. 183-188)

DIAGNOSIS. — Labial palpi dark apically. Frons very pale, almost white. Antennae pale, with a
single red line on dorsal surface of scape. Pronotal red stripe broad. All forewing crossveins very dark,

PENNY: LACEWINGS OF COSTA RICA 213

including gradates. Three to four inner and four outer gradate veins. Males with extremely large ab-
dominal spiracles. Male terminalia distinctive, with a knob at the apex of the gonapsis, large lateral
gonocristae, as well as long, dense gonosetae on the gonosaccus.

GEOGRAPHICAL DISTRIBUTION. — Adams and Penny (1987) record this species from the south-
ern U.S.A., Mexico, Cayman Islands, Cuba, Jamaica, Haiti, Puerto Rico, Barbados, Honduras, Vene-
zuela, Guyana, Surinam, and Brazil. Within Costa Rica it has been recorded from: ALAJUELA:
20 km S Upala; CARTAGO: Turrialba (C.A.T.IE.), GUANACASTE: 14 km S Canas.

ALTITUDINAL DISTRIBUTION. — Unknown.

ADULT FLIGHT PERIOD. — January to April, June, November, and December.

HABITAT PREFERENCE. — Adams and Penny (1987) mention this species being abundant on
hedgerows of native vegetation and occasionally citrus trees in Honduras.

Ceraeochrysa defreitasi Penny, new species
(Figs. 189-195)

HOLOTYPE. — Male, “Finca Las Cruces near San/ Vito, Puntarenas Province, Costa Rica,
1800 m elev./ 14.III.69 at black light/ Joe Sheldon #1969-142.” Holotype deposited in the Entomol-
ogy Collection of the California Academy of Sciences, San Francisco. Two male paratypes, same data
as holotype except 15.11.69, #1969-151 and #1969-154. One deposited at California Academy of Sci-
ences and other at Instituto de Biodiversidad, Santo Domingo de Heredia, Costa Rica.

DIAGNOSIS. — This is one of several Central American species of Ceraeochrysa with four dark
spots on the pronotum. However, unlike C. nigripedis, C. inbio, and C. elegans, there are no dark
markings on the meso- and metanota. The two species with only two dark spots (C. discolor has red
spots and a recurved ventral extension of the dorsal apodeme) on the pronotum (C. tauberae and C.
costaricensis) also have completely pale meso- and metanota, but C. tauberae has distinctly dark
swellings on the forewing at the apex of Cu and 2A. Ceraeochrysa costaricensis may be the most
closely-related species in that the male gonapsis has lateral arms, the dorsal apodeme of the ectoproct
has a ventral extension with apical point, the mediuncus has a pair of large caudolateral lobes and there
is an extensive field of gonocristae on the gonosaccus. However, the differences are also numerous: C.
defreitasi has four pronotal spots rather than two; the male gonapsis has only a single lateral arm; the
gonarcus has no mediodorsal plate; the mediuncus lobes are apically pointed, not truncate; and the
pair of dorsal lobes of the arcessus are better developed.

HEAD. — Frons, gena, clypeus, labrum, palpi and vertex yellow without dark markings.
Antennal scape, pedicel and flagellum yellow without dark markings.

THORAX. — Pronotum yellow with pair of dark reddish brown spots laterally on each side.
Meso- and metanota yellow without dark markings. Wings: Forewing length = 14.5 mm. Forewing
pale green without dark spots on membrane. Longitudinal veins pale green. Costal and radial
crossveins dark. Six to eight (six to seven in holotype) inner and eight to nine outer gradate crossveins
dark with dark margining of membrane. Inner gradate series sigmoidally curved while outer gradates
straight. Hind wing pale green without dark markings. Six inner and eight outer gradate crossveins.

ABDOMEN. — Pale yellow with a pair of small reddish spots posterolaterally on tergites three to
six. Male tergite 9 with small pointed dorsal lobe. Dorsal apodeme of ectoproct thick, with elongate
ventral projection with acute apical point below ventral margin of ectoproct. Gonapsis elongate with
subapical lateral lobe on left side and apical small dorsal point and more elongate ventral point.
Gonarcus with ovate lateral arms and no medial points or dorsal plate. Mediuncus with pair of small
dorsal points and more elongate pair of caudolateral lobes. Arcessus with pair of lateral acute points
and decurved terminal ventromedial point. Gonosaccus with extensive field of small gonocristae in
aligned rows and scattered lateral gonosetae.

214 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES
Volume 53, No. 12

ETYMOLOGY. — This species is named for Sergio de Freitas, a chrysopid systematist and ecolo-
gist who has done much to elucidate the fauna and biology of Brazilian agro-ecosystems.

Ceraeochrysa discolor (Navas, 1914)
(Figs. 196-201)

DIAGNOSIS. — This isa member of the cincta group, a group defined by a recurved branch of the
dorsal apodeme of the male ectoproct. It is the only member of this group witha single spot laterally on
each side of the pronotum. The only other member of this group with pronotal spots is C. inbio, which
has two dark spots on each side at the anterior and posterior margins. In addition to the pronotal spots,
there are at least two spots on the mesonotum. A double dorsal stripe on the antennal scape and a swol-
len 2a-3a crossvein also help to distinguish this species from all others. One of the scape stripes is
smaller than the other and the mesonotum spots may sometimes be faint. There are also small dark
spots under the dorsolateral margin of the pronotum and at the dorsolateral margin of the mesonotum
of some specimens.

GEOGRAPHICAL DISTRIBUTION. — This species was originally described from Volcan de
Chiriqui in Panama, within sight of the Costa Rican border. Within Costa Rica this species has been
collected at: ALAJUELA, Guanacaste National Park, San Ramon Field Station; GUANACASTE,
Pitilla Field Station, 9km S Santa Cecilia; Cacao Field Station, SW of Cacao Volcano;
PUNTARENAS, Monteverde Biological Reserve, San Luis; La Amistad National Park, Las Alturas
Field Station; and Las Cruces, 6 km S San Vito.

ALTITUDINAL DISTRIBUTION. — The Panamian record is from 760—1220 m. In Costa Rica from
620 to 1500 m.

ADULT FLIGHT PERIOD. — February, March, April, August, November, and December.

HABITAT PREFERENCE. — Label data indicate that this species 1s associated with forests, but this
genus is rarely collected in deep forest. It is probably a forest-edge dweller.

Ceraeochrysa elegans Penny, 1998
(Figs. 202—207)

DIAGNOSIS. — This is a very distinctive species. There are four dark red spots at the corners of
the pronotum, as well as a pair of red spots each on the meso- and metascuta. The abdomen has red
bands across the posterior margin of each segment. The apex of the posterior branch of Cu is swollen,
darkened and margined with dark on the membrane. In most of these respects it is similar to C.
nigripedis. However, C. elegans has a dark scape stripe and red bands on the abdomen, which are lack-
ing in C. nigripedis.

GEOGRAPHICAL DISTRIBUTION. — This species has been collected in northern Mexico and Costa
Rica. It can be assumed that it will be found in some intermediate areas as well. Within Costa Rica,
Ceraeochrysa elegans has been collected at. CARTAGO, Monumento Nacional Guayabo, A. C.
Amistad, G. Fonseca, 1 female (allotype) (INBIO); PUNTARENAS: Monteverde Biological Re-
serve, La Casona Station, E. Bello, 1 male (holotype) (INBIO); and Finca Las Cruces, 6 km S San
Vito de Java, J. E. Eger (TAMU).

ALTITUDINAL DISTRIBUTION. — 1100-1520 m.

ADULT FLIGHT PERIOD. — July, September, and December.

HABITAT PREFERENCE. — All three known localities for this species in Costa Rica have good
stands of primary forest, and at Monteverde this is almost the exclusive habitat. However, as this ge-
nus is not normally found deep within the forest, it probably dwells at the forest-edge.

PENNY: LACEWINGS OF COSTA RICA 21S

Ceraeochrysa everes (Banks, 1920)
(Figs. 208-214)

DIAGNOSIS. — This is a species with dark antennae and narrow, red pronotal stripes. The labial
palps are pale and there is a dark brown genal stripe, which will separate this species from C. claveri.
In the forewing only the costal, radial, and gradate crossveins are darkened. The male genitalia in-
clude a large medial hood on the gonarcus and large, dorsally-diverging entoprocesses.

GEOGRAPHICAL DISTRIBUTION. — The type is from French Guiana. It has been recorded under
various names from Cuba, Honduras, Panama, Colombia, Trinidad and Brazil. Within Costa Rica this
species has been recorded from: ALAJUELA, 20 km S Upala; CARTAGO, Turrialba (C.A.T.L.E.);
GUANACASTE, Guanacaste National Park; Maritza Field Station, near Orosi Volcano; 14 km S
Cafias; Finca La Pacifica, near Cafias; Rio Gongora, 4 km NE Quebrada Grande; and LIMON, Pan-
dora.

ALTITUDINAL DISTRIBUTION. — The Pandora locality is at about 100 m elevation, while the
Santa Rosa locality is at 300 m.

ADULT FLIGHT PERIOD. — February, June, and September.

HABITAT PREFERENCE. — The Pandora specimen was collected on understory vegetation near a
stream in primary forest. The La Pacifica adults were collected in a very similar situation, amongst
bushes along a river.

Ceraeochrysa gradata (Navas, 1913)
(Figs. 215—220)

DIAGNOSIS. — This species has pale frons, palpi and genae. The antennal scape bears a dorsal
dark stripe and faint indication of lateral stripe. The antennal flagellum is dark basally and pale api-
cally. The pronotum has a narrow, lateral red stripe. Superficially this species appears similar to C.
claveri, but this latter species has antennae darkened to the apex and a broader pronotal stripe.

GEOGRAPHICAL DISTRIBUTION. — Originally described from Guatemala. Within Costa Rica, it
has been collected at LIMON, Cahuita National Park; and PUNTARENAS, Monteverde.

ALTITUDINAL DISTRIBUTION. — Sea level to 1400 m.

ADULT FLIGHT PERIOD. — September.

HABITAT PREFERENCE. — Two females were collected amongst woody vegetation directly be-
hind the beach in very humid, swampy conditions, exposed to prevailing winds and sea spray.

REMARKS. — Ceraeochrysa indicata (Navas, 1914) and Ceraeochrysa infausta (Banks, 1946),
both recorded from Costa Rica, are new synonyms of Ceraeochrysa gradata (Navas, 1913).

Ceraeochrysa inbio Penny, 1997
(Figs. 221—226)

DIAGNOSIS. — This species is distinctive in having four dark spots on the pronotum and pair of
dark spots on the mesonotum. It lacks a pair of spots on the metascutum, which are present in C.
elegans and C. nigripedis. It also lacks the swollen apical veins of the posterior cubitus in the
forewing, which are present in the latter two species. The long ventral hook of the dorsal apodeme of
the male ectoproct indicate a close relationship with other members of the cincta group.

GEOGRAPHICAL DISTRIBUTION. — At present this species is known only from the type specimen
from Costa Rica: CARTAGO: La Amistad Biosphere Reserve, Guayabo National Monument.

ALTITUDINAL DISTRIBUTION. — 1100 m.

ADULT FLIGHT PERIOD. — July.

HABITAT PREFERENCE. — Unknown.

216 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES
Volume 53, No. 12

Ceraeochrysa josephina (Navas, 1926)

This species was originally described from Costa Rica. The original description contains no in-
formation that would allow identication of this species and the primary type cannot be located. Adams
(1982) synonymized Chrysopa angulata Navas, 1929 with Ceraeochrysa josephina without having
seen the type of the latter species. Ceraeochrysa angulata, from Colombia, has a distinctive swelling,
darkening and tiny crossvein at the fork of the forewing distal cubital cell, not mentioned in Navas’
original description of C. josephina. No specimens of Ceraeochrysa angulata have yet been recorded
from Costa Rica. Until such time as the type of C. josephina can be located or a neotype designated, I
think it best to consider Ceraeochrysa josephina as anomen dubium, and Ceraeochrysa angulata as a
valid species.

Ceraeochrysa nigripedis Penny, 1997
(Figs. 227-232)

DIAGNOSIS. — This species is very distinctive and should be immediately recognizable, even
when some markings are faded. There are always two dark spots on the mesoscutum and two more
black spots on the metascutum. Less frequently preserved are four red spots at the corners of the
pronotum, a lateral stripe on the antennal scape, and transverse yellow and red bands dorsally on each
abdominal segment. There is a darkened area of incrassate veins at the apex of CuP in the forewing.
Finally, the last tarsal segment of all legs is contrastingly dark brown to black. This is one of the most
colorful species of Ceraeochrysa.

GEOGRAPHICAL DISTRIBUTION. — So far, this species is known only from Costa Rica, recorded
from two specimens: PUNTARENAS, Monteverde Biological Reserve, La Casona Station, N.
Obando, | male, | female (INBIO, CAS).

ALTITUDINAL DISTRIBUTION. — 1520 m.

ADULT FLIGHT PERIOD. — November.

HABITAT PREFERENCE. — Unknown.

Ceraeochrysa pseudovaricosa Penny, 1998
(Figs. 233-238)

DIAGNOSIS. — The reticulated series of small, darkened cells in the middle of the inner gradate
series of the forewing appear very similar to that of Chrysopodes varicosa. However, the two species
can be distinguished by the lack of transverse bands on the frons of C. pseudovaricosa. The presence
of an elongate, spoon-shaped gonapsis in the male genitalia indicate its true affinities within
Ceraeochrysa.

GEOGRAPHICAL DISTRIBUTION. — So far, this species has only been collected at: Costa Rica:
PUNTARENAS: Parque Nacional La Amistad (8°57'N, 82°50'W).

ALTITUDINAL DISTRIBUTION. — 1500-1600 m.

ADULT FLIGHT PERIOD. — Mid-February to early March.

HABITAT PREFERENCE. — The nine known specimens of this species were collected in primary
forest by beating the understory vegetation near the edge of La Amistad National Park. This is an area
in the rain shadow of the Talamanca Mountains and in February receives very little rainfall. The for-
est, although evergreen, is very dry during this period.

PENNY: LACEWINGS OF COSTA RICA DAY

Ceraeochrysa sanchezi (Navas, 1924)
(Figs. 239-246)

DIAGNOsIS. — This is one of the species of Ceraeochrysa with pale palpi and dark antennae.
There is no stripe on the antennal scape, but rather, a darkened diffused reddish area on the lateral sur-
face. This lack of scape stripe separates it from C. claveri and C. indicata. It also has a yellow
mesonotum. The most distinctive features are found in the male terminalia: the elongate ventral pro-
jection of the ectoprocts encircling the abdominal apex, the double row of vertical gonocristae above
the apex of the ninth sternite, the flattened dorsal entoprocesses and the large, sclerotized teeth on the
gonosaccus.

GEOGRAPHICAL DISTRIBUTION. — This species was originally described from Cuba. Adams and
Penny (1987) reported collections from Mexico, Honduras and Brazil. Within Costa Rica, it has been
collected from ALAJUELA, 20 km S Upala; GUANACASTE, 8 km (5 mi) NE Canas; Finca La
Pacifica, near Canas; Finca Taboga, near Canas;14 km S Cafias; Guanacaste National Park; and
PUNTARENAS, Las Cruces, 6 km S San Vito.

ALTITUDINAL DISTRIBUTION. — Lowland areas around 300 m or less.

ADULT FLIGHT PERIOD. — December to March.

HABITAT PREFERENCE. — At La Pacifica adults were collected by beating bushes along a river
bank in February.

Ceraeochrysa smithi (Navas, 1914)
(Figs. 247-252)

DIAGNOSIS. — This is a striking species with black base to the antennal flagellum and broad, red,
lateral pronotal stripes. However, most distinctive are the red dorsal surface of the antennal scape and
broadly-triangular, red mark on the vertex at the base of each scape. The male also has a subapical dor-
sal projection on both sides of the ninth sternite.

GEOGRAPHICAL DISTRIBUTION. — This species is probably found throughout the Caribbean,
Central America, and northern South America. It was originally described from the Windward Islands
of the West Indies. When Adams (1982a) included the species in Ceraeochrysa he synonymized two
names under it: C. neotropica Navas, reported from Colombia and Brazil (Penny 1978) and C. poeyi
Navas from Cuba. Penny and colleagues (1997) reported C. smithi from U.S.A. (Florida). Additional
specimens have been reported from Mexico (as far north as Baja California Sur), Guatemala, Hondu-
ras, Costa Rica, and Panama. Within Costa Rica specimens are known from: CARTAGO, Turrialba;
GUANACASTE, Guanacaste National Park, Agua Buena, Cacao Field Station on the southwest side
of Cacao Volcano; Finca Jenny, 31 km N Liberia; Finca La Pacifica, near Canas; and Rio Gongora,
6 km northeast of Quebrada Grande de Liberia; and HEREDIA, Puerto Viejo.

ALTITUDINAL DISTRIBUTION. — From 220 m at Agua Buena to 1000—1400 m at the Cacao Field
Station.

ADULT FLIGHT PERIOD. — February, August, September, and November-December.

HABITAT PREFERENCE. — Adults have been collected in bushes near Rio Corobici at La Pacifica
in February.

Ceraeochrysa tauberae Penny, 1997
(Figs. 253-258)

DIAGNOSIS. — Ceraeochrysa tauberae is one of the Central American species with two dark
spots on the pronotum. This species has darkened, incrassate veins at the apex of CuP in the forewing
and forms an interesting convergence with C. nigripedis, which has two black spots on the

218 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES
Volume 53, No. 12

metascutellum. However, C. tauberae has a pale metascutellum and a darkened, incrassate a2-a3
crossvein, which at rest is folded over the metascutellum. The inner and outer gradate veins are
heavily margined with darkened membrane.
GEOGRAPHICAL DISTRIBUTION. — This species has only been collected in Costa Rica:
CARTAGO, near Eslabon; and PUNTARENAS, Finca Las Cruces, 6 km S San Vito.
ALTITUDINAL DISTRIBUTION. — The Las Cruces locality 1s between 1000-1400 m elevation.
ADULT FLIGHT PERIOD. — Late September to early October and December.
HABITAT PREFERENCE. — The Finca Las Cruces locality is the Wilson Botanical Garden and as-
sociated undisturbed forest.

Ceraeochrysa valida (Banks, 1895)
(Figs. 259-264)

DIAGNOSIS. — In live material the frons is pale yellow to white, whereas other species have a
pale frons only in faded, preserved material. The dark tips of labial palpi are unusual for this genus.
Antennae are pale, with a very broad, dorsal, red scape stripe. The lateral red pronotal stripe is very
broad and short There are also only one to three inner gradate veins in the forewing. The male
ectoprocts are ventrally extended at their apices.

GEOGRAPHICAL DISTRIBUTION. — This species is broadly distributed from the southeastern
U.S.A. south to Ecuador and Brazil. Within Costa Rica it has been found at: GUANACASTE, 14 km
S Cafias; Finca La Pacifica, near Cafias; Finca Taboga, near Cafias; and SAN JOSE, Escazu.

ALTITUDINAL DISTRIBUTION. — Lowland areas at about 300 m near Canas, to about 1000 m in
the central valley near San José.

ADULT FLIGHT PERIOD. — January to March, May, September, October, and December, but
most common from January to March..

HABITAT PREFERENCE. — At Canas adults have been collected by beating bushes near the Rio
Corobici in February.

Chrysoperla Steinmann, 1964

This genus is probably the most common group of green lacewings worldwide, partly because of
its preference for open agricultural and pasture lands, where adults feed on grass pollen. Several spe-
cies in this genus have been mass reared for release in fields and orchards as augmentative biological
control agents. They are among our most important beneficial insects (Tauber et al. 2000) and have
great potential for control of white flies, aphids, scale insects and mealybugs in Costa Rica. Eggs are
laid singly near concentrations of honeydew excreted by homopterous insects. However, larvae feed
on a wide variety of insects which they glean from foliage as they search the plants. Larvae do not
cover their bodies with debris or the bodies of their prey. Adults can often be found in large numbers in
pastures, lawns, and other areas where grasses are plentiful. All New World species are pale green
with a red spot on the gena below the eyes. There is a medial yellow stripe along the mid-dorsal line of
the thorax and abdomen. Some species in temperate climates overwinter as hibernating adults and
body coloration changes to reflect a changed physiology. As colder weather approaches, body color-
ation becomes pale brown and a pair of red spots appear on each segment of the abdomen. Adults mi-
grate in very large numbers to the forest, where they rest under live leaves in milder climates and
amongst dead, fallen leaves in harsher, colder climates. The genus is worldwide, with 50 known spe-
cies. Only two species are known from Costa Rica.

PENNY: LACEWINGS OF COSTA RICA 219

KEY TO COSTA RICAN CHRYSOPERLA

IMA eitexawitha pair ofred submedial spots, often fused medially .% 3.4.5. 2s. eee ee eae C. exotera
Meniexecntinclynale creenitOypal esVCllOW soc A i Gs see, Seti BGS cp Ss cig tieisp rem a upd swan cb cata ee- C. externa

Chrysoperla exotera (Navas, 1913)
(Figs. 265—267)

DIAGNOSIS. — This species is most easily recognized by the large reddish submedial to medial
marks in the center of the vertex. The male genitalia are also diagnostic, i.e., male tignum evenly ta-
pered and strongly arched.

GEOGRAPHICAL DISTRIBUTION. — This species is known from Mexico, Guatemala, Honduras,
and Costa Rica. Within Costa Rica it has been collected at, GUANACASTE: Finca La Pacifica; and
SAN JOSE, Escazu.

ALTITUDINAL DISTRIBUTION. — Pacific lowland areas below 200 m to mid-elevations around
1000 m.

ADULT FLIGHT PERIOD. — February to June.

HABITAT PREFERENCE. — Like other members of the genus, this species prefers open, grassy ar-
eas, such as the suburban, residential area of Escazu and near disturbed habitats along the Rio
Corobici at Finca La Pacifica.

Chrysoperla externa (Hagen, 1861)
(Figs. 268-270)

DIAGNOSIS. — Characteristics include a pale green vertex with no red markings and a male
tignum that is swollen at mid-length and has very little arch to it.

GEOGRAPHICAL DISTRIBUTION. — This is the most widespread species of Chrysoperla in the
New World, ranging from U.S.A. (Florida) south to Argentina, and can probably be found in all coun-
tries in between. It has also been collected on Cocos Island (Costa Rica) and Easter Island. Wherever it
occurs, it is also among the most abundant species present, and is the Neotropical ecological counter-
part of C. plorabunda and C. rufilabris in North America and C. carnea in Europe. Within Costa Rica
it can probably be found anywhere where grasslands exist below 2200 m. Records include:
ALAJUELA, 20km south of Upala; GUANACASTE, Santa Rosa National Park, Quebrada
Pedregal; 14 km south of Canfas; 6 miles north of Canas, Finca La Pacifica, near Canas; LIMON,
Hitoy-Cerere Biological Reserve; PUNTARENAS, La Amistad National Park, Las Alturas; La
Amistad National Park, upper Rio Bellavista; Osa Peninsula, Rincon; and SAN JOSE, Escazt.

ALTITUDINAL DISTRIBUTION. — The highest elevation known is 2200 m. Below this level,
adults can be found almost anywhere where grasslands exist.

ADULT FLIGHT PERIOD. — Records indicate January through April, although adults probably
occur throughout the year.

HABITAT PREFERENCE. — Adults feed on grass pollen and are frequently very abundant in grass-
lands. They are also taken in orchards and on cultivated crops in Nicaragua. They are attracted to lights
in appropriate areas.

Chrysopodes Navas, 1913
Chrysopodes is a Neotropical genus with 32 recognized species (Brooks and Barnard 1990). The

genus is most easily defined by the male genital components. The tignum is absent, the gonapsis usu-
ally absent, with the gonarcus complex often medially inflated and bulbous. Chrysopodes is usually

220 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES
Volume 53, No. 12

divided into two subgenera with distinctly different appearances. The subgenus Neosuarius has nar-
rower forewings, particularly in the costal area. The mandibles are stout, blunt-tipped and asymmetri-
cal. The basic body coloration is often dark, particularly in southern South American species. The
subgenus Chrysopodes has broader wings with an expanded costal area. They appear to be fragile fli-
ers with very long setae on veins of the forewing. Mandibles are narrow, sickle-shaped and symmetri-
cal. The basic body coloration is always pale green, changing to straw yellow or pale brown after
death. Despite the general differences between the subgenera, some species of Neosuarius have pale
green coloration, and the basic structures of male and female genitalia are quite similar. The two
groups are clearly closely related and were grouped into a single genus long before recognition of its
subgenera. Several of the Costa Rican species in this genus have very irregular or swollen venation in
the middle of the forewing with corresponding dark coloration, creating a focal point of visual atten-
tion. Both subgenera appear to reach maximum species richness in South America. Eight species of
Chrysopodes are currently known from Costa Rica and more are expected to be found.

KEY TO COSTA RICAN CHRYSOPODES

1. Forewing with a small, dark, reticulate network of veinlets at middle of inner gradate series .... . C. (C.) varicosa
1’. Forewing with gradate series forming a stepwise series, with an occasional auxillary crossvein, but without a small
reticulate network: of veinlets:2)s, 2); ears sys crepe sl sles eee bo leeds ays 5S) 2
2(1). Forewing intramedian vein reaching terminal crossvein, forming a quadrate cell ................... 3
2'. Forewing intramedian vein not reaching terminal crossvein, forming an ovate intramediancell ........... 4

3(2). Antennal scape with a dorsal stripe; frons with a dark transverse band; vertex with a V-shaped dark mark posterior to

ANteMMAlDASESs. cncex, ec Morse: Ee Meiuln GeO Saye Weak ree es aes: cian OM ee C. (N.) oswaldi
3’. Antennal scape entirely pale; frons dark laterally, pale medially; vertex pale except for a small dark mark at
Gaudolateral manginray 2s 5 eo ahead eee Ck hg ese Goel meeps Saye se oe re C. (C.) nevermanni

4(2) Forewing with a darkened spot at fourth anal crossvein along hind margin; forewing inner gradate series heavily

marcinedwwithidarkenedimenibrane) yea. ei icons diciete) ee 2 ec ee C. (C.) costalis
4'. Forewing without dark spots along hind margin; forewing inner gradate series not margined, membrane entirely

thanS parent 2 os, Oty 4 Spee eo bee ee ee eee ee yh eS oe le ch ot ae 5
5(4). Pronotum with four discrete dark red spots; male forewing Rs swollen .............. C. (N.) crassinervis
5’. Pronotum either pale, or with red markings at anterior margin, or with longitudinal red stripes, but not four discrete

red\spotssmale forewineiRs swollenionnot. 7. =). 365 3 cee = 3) ees ee = tone cee ee 6

6(5). Frons entirely pale; maxillary palpi dark only at apex; pronotum with longitudinal red stripe; male forewing Rs not

SWOMEM PEAS. gene eerh ce toes pe Tite Petes bel bein, Mined oh Me Ue eve eens at eer rs 7 a C. (C.) victoriae
6’. Frons with two transverse red bands below antennal bases; maxillary palpi dark basally or entirely pale; pronotum
pale or with dark red markings only at anterior margin; male forewing Rs at least somewhat swollen ........ 7

7(6). Heavy-bodied, with pronotum pale, wider than long; maxillary palpi entirely pale; gradate crossveins green,

unmarcined::, S88. oa Bee ee ae ae ee or oes Geena a aes nN Pao fb C. (N.) collaris
7’. More slender-bodied, with pronotum longer than wide, with dark red spots at anterior margin; maxillary palpi dark,
except pale at apex; gradate crossveins dark with dark margining ................. C. (N.) crassipennis

Chrysopodes (Chrysopodes) Navas, 1913

The characters given above in the generic treatement will serve to distinguish this subgenus. The
most recent revision of Chrysopidae (Brooks and Barnard 1990) included 21 species in this subgenus.
They are distributed from Guatemala south to Argentina, with greatest species richness apparently oc-
curring in the Amazon Region and other parts of northern South America. Four species are known
from Costa Rica.

PENNY: LACEWINGS OF COSTA RICA 221

Chrysopodes (Chrysopodes) costalis (Schneider, 1851)
(Figs. 292-294)

DIAGNOSIS. — This species has the broad forewing costal area typical of most Chrysopodes
(Chrysopodes), but can be distinguished from other species by the heavier dark markings of the wing.
The inner gradate series is heavily infuscated, and the distal part of the anal margin has apical vein
twiggings darkened, resulting in the appearance of a stripe along this margin. However, the most dis-
tinguishing feature of the forewing is a single crossvein on the hind margin about halfway to the apex
that is dark, swollen, and heavily infuscated in the adjacent membrane. The Amazonian species
Chrysopodes (C.) duckei and C. (C.) pulchella also have the distinctive small dark area along the anal
margin, but the spot is much smaller in these species and they have heavy dark margining of the base
of the Rs forks, which 1s lacking in C. costalis.

GEOGRAPHICAL DISTRIBUTION. — The original description only indicates “America” as the lo-
cality. No additional published distributional records are known. Within Costa Rica, this species has
been collected at: LIMON: Pandora, Valle de Estrella.

ALTITUDINAL DISTRIBUTION. — The Pandora locality is at the base of a small hill at the edge of
“Star Valley” at less than 200 m elevation.

ADULT FLIGHT PERIOD. — September.

HABITAT PREFERENCE. — The single known Costa Rican individual was collected resting be-
neath the leaves of young trees, along a small stream. The area was heavily shaded by primary
overstory forest and very humid from its proximity to the stream.

Chrysopodes (Chrysopodes) nevermanni Navas, 1928
(Figs. 295-297)

DIAGNOSIS. — This species is most easily distinguished by its quadrate intramedian cell. This
and related species with quadrate intramedian cells also have dark palpi and a dark base to the
branches of the forewing radial sector, except C. costalis. The two Brazilian species in this group, C.
(C.) duckei and C. (C.) pulchella have an irregular third gradate series and a dark spot at one crossvein
about halfway to the apex along the hind margin. The other member of this group, C. (C.) inornata
from Colombia, lacks this forewing spot, but has lateral red stripes on the pronotum, which are lacking
in C. (C.) nevermanni.

GEOGRAPHICAL DISTRIBUTION. — This species was originally descibed from the plains between
Limon and La Mercedes on the Caribbean side of the Talamanca Mountains. Other records are from
Pandora and Hitoy-Cerere Biological Reserve in Valle de Estrella of LIMON Province. So far this
species is known only from the coastal plain of southeastern Limon Province.

ALTITUDINAL DISTRIBUTION. — From the margin of the “Star Valley” at under 200 m elevation
and near Limon at 150-300 m.

ADULT FLIGHT PERIOD. — March and September.

HABITAT PREFERENCE. — Two individuals were collected resting beneath the leaves of young
trees, along a small stream. The area was heavily shaded by primary overstory forest and very humid
from its proximity to the stream.

Chrysopodes (Chrysopodes) varicosa Navas, 1914
(Figs. 298-303)

DIAGNOSIS. — This species is very distinctive because of the small cluster of veinlets in the mid-
dle of the inner gradate series of the forewing, which 1s accentuated by the darkening of the membrane
surrounding these veinlets. The only other species with this distinctive feature is Ceraeochrysa

222 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES
Volume 53, No. 12

pseudovaricosa which, in addition to its Ceraeochrysa-type male genitalia with an elongate gonapsis,
also has a lateral stripe on the antennal scape, darker brown markings on the frons, and no transverse
red stripes below the antennal bases on the frons.

GEOGRAPHICAL DISTRIBUTION. — This species is known from Guatemala (type specimen) and
Costa Rica. In Costa Rica, it is known from: ALAJUELA: 20 km S Upala; Quebrada Virgencita,
10.2 km S Bajos del Toro (10.168°N, 84.326°W); and PUNTARENAS: Monteverde Biological Re-
serve; Las Alturas; Fila Cedro.

ALTITUDINAL DISTRIBUTION. — This species appears to prefer mid-elevations from
1500-2000 m, although the site at Upala must be lower than this. The Guatemalan type is from 400 m
elevation.

ADULT FLIGHT PERIOD. — February, July, and September.

HABITAT PREFERENCE. — Known collecting sites are in tall primary forest.

Chrysopodes (Chrysopodes) victoriae Penny, 1998
(Figs. 304-309)

DIAGNOSIS. — This is the only species of Chrysopodes yet known from Costa Rica with dark an-
tennae (at least basally). The red pronotal stripes coupled with lack of markings on the frons is also
distinctive, as is the trilobate apex to male ninth sternite.

GEOGRAPHICAL DISTRIBUTION. — This species is known only from the holotype male collected
at: LIMON: Pandora, Valle de Estrella.

ALTITUDINAL DISTRIBUTION. — Less than 200 m.

ADULT FLIGHT PERIOD. — September.

HABITAT PREFERENCE. — The holotype was collected resting beneath the leaves of young trees,
along a small stream. The area was heavily shaded by primary overstory forest and very humid from
its proximity to the stream.

Chrysopodes (Neosuarius) Adams and Penny, 1987

The characters mentioned in the generic key and discussion above will delimit this subgenus.
Brooks and Barnard (1990) listed 11 species, which range from southeastern U.S.A. and Mexico
south to Chile and Argentina. One species, Chrysopodes (Neosuarius) nigripilosa is found only on the
Galapagos Islands. Five species are known only from Chile and austral South America, and this area
appears to be the most species rich. Two species have broad distributions. Chrysopodes (Neosuarius)
divisa is known from Jamaica and Cuba south to Argentina, while C. (N.) collaris is known from
southeastern U.S.A. and Mexico (Baja California) south to Guyana and Venezuela. All other species
appear to have much more restricted distributions. The four described species from Costa Rica are
treated below. There is, in addition, one undescribed, apparently new, species from Costa Rica char-
acterized by heavy infuscation of the membrane surrounding crossveins, which is reminiscent of C.
(N.) limbata and C. (N.) parishi, but has a dark lateral scape stripe. Five females of this form have been
studied, however, until a male is found, this species shall remain unnamed.

Chrysopodes (Neosuarius) collaris (Schneider, 1851)
(Figs. 271-276)

DIAGNOsIS. — This is one of the more distinctive species of green lacewings. It is a rather
thick-bodied species with pronotum broader than long. It superficially resembles species of Nacarina
in the Belonopterygini, although the antennae are thinner. There is a distinct complete double trans-
verse red line below the antennae on the frons, which at times becomes a pink suffusion. In the male

PENNY: LACEWINGS OF COSTA RICA 223

genitalia the ectoproctal apodeme forks at the callus cercus and the ventral fork extends
ventrocaudally as an incurved apical projection. The male arcessus has a thick internal brace, which is
lacking in the related C. (N.) divisa.

GEOGRAPHICAL DISTRIBUTION. — This is a wide-ranging species known from the U.S.A.
(Florida and Texas), and Mexico (Baja California) south through Central America and the Antilles to
Colombia, Venezuela, and Guyana. Within Costa Rica specimens have been recorded from:
ALAJUELA, 20 km S Upala; CARTAGO, Playa del Coco; GUANACASTE, Santa Rosa National
Park, Rio Poza Salada; Quebrada Guacimo; Quebrada Pedregal; 14 km S Canas; Finca La Pacifica,
near Canas; and PUNTARENAS, Boca de Barranca.

ALTITUDINAL DISTRIBUTION. — 10-300 m.

ADULT FLIGHT PERIOD. — February, May, June, July, and September.

HABITAT PREFERENCE. — At Finca La Pacifica this species was collected at lights near the Rio
Corobici.

Chrysopodes (Neosuarius) crassinervis Penny, 1998
(Figs. 277—282)

DIAGNOSIS. — This species is immediately recognizable from its two pair of dark spots on the
pronotum and the heavily swollen forewing veins in the male. No other species of Chrysopodes from
Costa Rica is known to have four pronotal spots, although this feature is fairly common among species
of Ceraeochrysa and Plesiochrysa. Another Costa Rican species of Chrysopodes, in which males
have swollen forewing veins, is C. crassipennis, but this species lacks the four pronotal spots.

GEOGRAPHICAL DISTRIBUTION. — So far, this species has only been collected in Costa Rica.
Known localities are: GUANACASTE: Rio Orosi(10.991°N, 85.928°W): LIMON: Parque Nacional
Tortuguero, Rio Tortuguero, 3.5 km S Tortuguero (10.509°N, 83.504°W); and PUNTARENAS:
Finca Las Cruces, 6 km S San Vito de Java; Rio Jaba at rock quarry, 1.4 km W Las Cruces (8.79°N,
82.97°W), 16 km (10 m1) S of Rincon de Osa.

ALTITUDINAL DISTRIBUTION. — Sea level to 1280 m.

ADULT FLIGHT PERIOD. — February, March, April, August, and September.

HABITAT PREFERENCE. — At Las Cruces this species was collected at an ultraviolet light. The
area contains both a botanical garden and primary forest.

Chrysopodes (Neosuarius) crassipennis Penny, 2000
(Figs. 283-285)

DIAGNOSIS. — The two most distinctive characteristics of this species are the darkly margined
gradate crossveins and swollen (incrassate), forewing, radial sector veins of males.

GEOGRAPHICAL DISTRIBUTION. — This species appears to be rather common ina very restricted
area, all within Costa Rica. Although a single female has been collected 14km S Canas
(GUANACASTE) where a considerable amount of collecting has been done, all other known speci-
mens (more than 100) are from GUANACASTE, 3 km SE Rio Naranjo.

ALTITUDINAL DISTRIBUTION. — The two known localities both lie below 200 m.

ADULT FLIGHT PERIOD. — Virtually every month of the year.

HABITAT PREFERENCE. — Unknown.

224 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES
Volume 53, No. 12

Chrysopodes (Neosuarius) oswaldi Penny, new species
(Figs. 286-291)

HOLOTYPE. — Male from “10 mi. S. Rincon de Osa / Puntarenas Proince, Costa / Rica,
4—12.]11.69/ J. K. Sheldon Field #/.” The holotype is deposited in the Entomology Collection of the
California Academy of Sciences, San Francisco. Damage: Right foreleg and hindleg missing. Abdo-
men detached and placed in genital vial below specimen. Left forewing and hind wing detached and
placed on card below specimen.

HEAD. — Frons pale yellow with dark red transverse band below antennal bases; not extended to
lateral margin. Gena dark red. Clypeus pale yellow medially, dark red laterally. Labrum pale yellow.
Maxillary palp segments dark basally, pale apically. Labial palps pale. Vertex pale yellow with dark
red V-shaped mark posterior to antennal bases and small dark spot medially at posterior margin.
Antennal scape pale yellow with dorsal longitudinal dark red stripe. Pedicel and flagellum pale yel-
low.

THORAX. — Pro-, meso- and metanota pale yellow, without markings. Pleural areas and legs
pale yellow without markings. Wings: Forewing length = 15.5 mm. Forewing pale yellow without
dark markings on membrane. Longitudinal veins pale, crossveins dark. Five to six inner and nine
outer dark gradate crossveins. Hind wing entirely pale; apex acutely pointed. Five inner and seven to
eight outer pale gradate crossveins.

ABDOMEN. — Pale green without markings. Sternites without microtholi. Male ninth sternite
evenly tapered to rounded apex. Dorsal apodeme of ectoproct with two ventral extensions: proximal
extension apically truncate; distal extension medially curved to acute point. Tignum and gonapsis ab-
sent. Gonarcus dorso-ventrally flattened; lateral arms simple quadrate plates. Arcessus bulbous with
apically decurved point.

RELATIONSHIPS. — The double ventral projection of the dorsal apodeme of the ectoproct is
found in two other species of Chrysopodes (Neosuarius): C. collaris and C. divisa. Several characters
clearly separate these species. Chrysopodes (N.) collaris isamuch more robust species with pronotum
wider than long, and forewing venation entirely green. Both C. collaris and C. divisa have a double
transverse band on the frons (Fig. 272) and both species lack the V-shaped mark posterior to the
antennal base and the dorsal scape stripe. The intramedian cell is quadrangular in C. (N.) oswaldi, but
this character state is somewhat variable, and since there is but one known specimen, this character
must, for now, be used with some caution.

ETYMOLOGY. — This species is named for John D. Oswald, who has done much to advance our
knowledge of neuropteran systematics and phylogeny.

Meleoma Fitch, 1855

This genus is closely related to Chryvsopa but most species have one very distinctive fea-
ture—males have simple to elaborate structures between the antennae and on the frons. In different
species these may range from a simple protuberance to a deep pit with opposing, setiferous prolonga-
tions. The antennal scape is very long and arched in some males or flattened and concave in others. A
few species lack these special head structures. Species of Meleoma are always rather large and robust.
This isa New World genus with a center of extant diversity in the southwestern United States and cen-
tral Mexico. There are 25 known species distributed from southern Canada south to Colombia and
Venezuela. Only two species are known from Costa Rica. Both have distinctive red stripes along the
margins of the eyes, on the vertex and along the genae to the mandibular bases. Some forewing
crossveins are red, slowly changing to black after death.

PENNY: LACEWINGS OF COSTA RICA 225

KEY TO COSTA RICAN MELEOMA

1. Forewings with only seven or eight outer gradate crossveins; male with elongate medial process between antennal
bases, with two apical tufts of hairs and lateral depression between antennal bases and eyes; male mediuncus pointed
1 aa gM GES i a0, ROMER licanes ORRIN Re span Pe ee cee ar eae eT M. titschaki

I’. Forewing with ten or more outer gradate crossveins; male with only a tiny, rounded projection between antennal
bases, and no depression between antennal bases and eyes; male mediuncus expanded and rounded apically M. poolei

Meleoma poolei Adams, 1969
Begs. 310, 313, 315)

DIAGNOSIS. — This species has more gradate veins than M. titschaki. This character may be used
to separate females. Males have a very poorly-developed medial horn between the antennal bases, and
no pits lateral to the antennal bases. The male genitalia has an elongate mediuncus which terminates in
a spoon-shaped process.

GEOGRAPHICAL DISTRIBUTION. — This species was described from Rancho Grande in Vene-
zuela, and until now was only known from that locality. Within Costa Rica this species has been col-
lected at! CARTAGO, Embalse El Llano; and SAN JOSE, El Salvage, Rio Tabarcia, 8 km (road) E
Palmichal, 9.847°N, 84.164°W.

ALTITUDINAL DISTRIBUTION. — 1650 m.

ADULT FLIGHT PERIOD. — January and September.

HABITAT PREFERENCE. — The Embalse El Llano locality is primary forest, with only mimimal
disturbance, but as specimens were collected with light traps, individuals may have been drawn from
neighboring cultivated areas.

Meleoma titschaki Navas, 1928
(Figs. 311, 312, 314)

DIAGNOSIS. — This species has fewer gradate veins than M. poolei. Males have an elongate pro-
jection between the eyes which terminates in two downwardly oriented tufts of hair. There is also a
shallow pit lateral to each scape. The male mediuncus has a recurved, acutely pointed apex.

GEOGRAPHICAL DISTRIBUTION. — Originally described from Costa Rica, but neotype desig-
nated from southern Mexico. Within Costa Rica this species is known from: ALAJUELA, San Ramon
Biological Reserve, Rio San Lorenzo; Cerro Campana, 6 km NW Dos Rios; Rio Peje, | km SE San
Vicente; GUANACASTE, SW side of Cacao Vulcano, Cacao Biological Station; PUNTARENAS,
Rio Cotén, Las Tablas; Monteverde; and SAN JOSE, San José (original type); Rio Tabarcia, 8 km E
Palmichal.

ALTITUDINAL DISTRIBUTION. — 640-1990 m. Most records are from above 1400 m.

ADULT FLIGHT PERIOD. — January to September, with one record from November—December,
perhaps late in December.

HABITAT PREFERENCE. — Unknown.

Plesiochrysa Adams, 1982b

Plesiochrysa was originally described as a subgenus of Chrysopa and indeed they are closely re-
lated, with males with much the same genital elements present. However, Chrysopa as currently de-
fined does not exist in the Neotropical Region. Plesiochrysa now has 22 recognized species from
southern Asia, Australia, Oceania and the New World tropics (Brooks and Barnard 1990). Two spe-
cies are found in Costa Rica, and are recognizable by their robust size and four red spots on the

226 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES
Volume 53, No. 12

pronotum, similar to North American Chrysopa quadripunctata. The male genital structures of the
two Costa Rican species are indistinguishable.

KEY TO COSTA RICAN PLESIOCHRYSA

1. Pronotum wider than long; forewing with gradate series divergent. .................... P. brasiliensis
1’. Pronotum longer than wide; forewing with gradate series parallel. ...................... P. elongata

Plesiochrysa brasiliensis (Schneider, 1851)
(Figs. 316-318)

DIAGNOSIS. — This species can be quickly separated from P. elongata by the distinctly diver-
gent series of gradate crossveins in the forewing. The pronotum is also somewhat wider than in the lat-
ter species.

GEOGRAPHICAL DISTRIBUTION. — This is a wide ranging species of chrysopid, collected in
Cuba, Mexico, Honduras, Nicaragua, Costa Rica, Panama, Colombia, Trinidad, Guiana, Ecuador and
Brazil. Within Costa Rica, records are from: ALAJUELA, Cerro Campana, 6 km NW Dos Rios;
GUANACASTE, Santa Rosa National Park (SRNP), Maritza Biological Station, Tempisque; SRNP,
Rio San Josecito; SRNP, Quebrada Pedregal; Rio Gongora, 4 km NE Quebrada Grande; 4 km N
Cafias; HEREDIA, Chilamate; Braulio-Carrillo National Park, El Ceibo Biological Station; LIMON,
Tortuguero National Park, 3.5 km S Tortuguero; PUNTARENAS, near Villa Neily; La Amistad Na-
tional Park, Las Tablas Protective Zone, Sitio Coton; Las Cruces, near San Vito; and SAN JOSE,
Finca La Caja (La Uruca); San José.

ALTITUDINAL DISTRIBUTION. — From sea level at Santa Rosa National Park to 1800 m near Las
Cruces.

ADULT FLIGHT PERIOD. — Present throughout the year, with the most numerous collections be-
tween April and July.

HABITAT PREFERENCE. — This species is attracted to lights and has been collected along large
rivers.

Plesiochrysa elongata (Navas, 1913)
(Figs. 319-321)

DIAGNOSIS. — This species is similar to P. brasiliensis except that the gradate series of
crossveins in the forewing are parallel and the pronotum is longer than wide.

GEOGRAPHICAL DISTRIBUTION. — It is known from Jamaica, Mexico, Cayman Islands, Hondu-
ras, Nicaragua, Costa Rica, Panama, Guadaloupe, Peru, and Brazil. Within Costa Rica records are
from: CARTAGO, Turrialba, Santa Teresito de Lajas; GUANACASTE, Santa Rosa National Park
(SRNP), Quebrada Guacimo; SRNP, Rio San Josecito; Rincon de la Vieja National Park, Rio Negro;
west side of Orosi Vulcano, Maritza Biological Station; PUNTARENAS, Monteverde; and SAN
JOSE, San José; 96 km S San José, Hotel La Georgina.

ALTITUDINAL DISTRIBUTION. — 255-3100 m (the latter at Hotel La Georgina).

ADULT FLIGHT PERIOD. — Most months between March and November.

HABITAT PREFERENCE. — This species has been collected at lights in disturbed, cultivated habi-
tats.

REMARKS. — This species was reported from Costa Rica under the name Chrysopa angusta
(Navas, 1914), now considered a synonym of Plesiochrysa elongata.

PENNY: LACEWINGS OF COSTA RICA 227

Ungla Navas, 1914

This genus was originally described for a composite insect consisting of a chrysopid clutching the
leg of an ant-lion (see Adams 1975). Despite its composite nature, the name remains available, and
Brooks and Barnard (1990) chose the chrysopid part of the composite as the original type ofa valid ge-
nus. The distinguishing characters of this genus are mostly found in the male terminalia. The male
ectoproct is short and not tapered basally. The dorsal apodeme, when present, is vertically oriented
through the callus cercus. The mediuncus forms a single elongate point, unlike the shorter, bulbous
mediuncus of Chrysopodes. Brooks and Barnard (1990) recognized four species from Argentina and
Peru. However, this genus is now known from Costa Rica and other undescribed species are known
from intervening areas. Only one species is known from Costa Rica.

Ungla pallescens Penny, 1998
(Figs. 322-327)

DIAGNOSIS. — This species can be separated from all others by the generic characters, particu-
larly the short, untapered male ectoproct with vertical dorsal apodeme. It is also the only chrysopid
known from Costa Rica which has mottled brown coloration imposed on blue-green tints while alive.
The widely dispersed whitish highlights when dead also are characteristic, and hence the name
pallescens, Latin for pale.

GEOGRAPHICAL DISTRIBUTION. — At this time, this species is only known from: Costa Rica:
PUNTARENAS, Monteverde, Pension Quetzal.

ALTITUDINAL DISTRIBUTION. — Collected at about 1500 m.

ADULT FLIGHT PERIOD. — September.

HABITAT PREFERENCE. — All three known specimens were collected at a porch light overlook-
ing a disturbed pasture and plantation habitat. The immediate surroundings contained native trees in-
terspersed with bananas, fruit trees, ornamental shrubs, and a grassy yard in a fairly open
environment.

NOTE. — Two additional chrysopids have been recorded from Costa Rica: Chrysopa bulbosa
(Navas, 1926) and Chrysopa sarta Banks, 1914. Neither species description contains enough useful
information to allow proper placement of these species and at this time must be considered incertae
sedis.

FAMILY CONIOPTERYGIDAE

by
Martin Meinander

Coniopterygids, or dustywings, are apparently rather abundant in Costa Rica, but are often over-
looked because of their small size. The dustywings are quite distinctive, small creatures, less than
3 mm long. In general appearance the adults bear a considerable resemblance to the white-flies
(Homoptera: Aleyrodidae), with a white waxy substance covering the whole insect. Larvae are arbo-
real, highly predatory, feeding on aphids (Homoptera: Aphidiidae) and scale insects (Homoptera:
Coccoidea). Adults are most active at dusk and dawn, when they sometimes can be seen in swarms.
Accurate identification of the species in this family generally requires an examination of the male gen-
italia, while identification to generic level can be done based on wing venation. Identification of fe-
males is presently impossible for many species, although in the future female genitalia will certainly
yield accurate key characters. The primary difficulty in using female genitalia is weak sclerotization
of important structures.

228 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES
Volume 53, No. 12

The various species are easily beaten from trees and bushes and are attracted to light.
In addition to the species recorded from Costa Rica, species recorded from Nicaragua and Pan-
ama are also included, as many species from the latter countries probably also occur in Costa Rica.

KEY TO THE SUBFAMILIES AND GENERA
OF COSTA RICAN CONIOPTERYGIDAE

1. Head capsule in lateral view distinctly higher than broad, genae broad (Fig. 329); two radio-medial crossveins (Figs.
331-332: r-m) in middle of forewing; in hind wing, Rs branching off from R very near base of wing (Figs. 331-332)
Ba as tase Gi eens lend Aarne Subfamily Aleuroptenyeinae’= .- 2. 2 se)

1’. Head capsule in lateral view fairly spherical, genae narrow (Fig. 330); one radio-medial crossvein in middle of
forewing (Figs. 333-334: r-m); in hind wing Rs not branching off from R very near base of wing (Figs. 333-334). ...

ASR CEEEO Ee hue oe: ety Bee: Subfamily Conioptenyeinae’s 2 = 54.2%.) 5 2s ee eS
2(1). Radial crossvein in hind wing (Fig. 331: r-r) striking Rs on stemoffork ................. Aleuropteryx
2'. Radial crossvein in hind wing (Fig. 332: r-r) striking Rs on branchR,,,................... Neoconis
3) sMrofhind wine mmtorkedi(Big9333))— aie sess Goer eee fe colts esc ee eke recta ee Coniopteryx
3). M.of hindwing forkedintoM 45 and M,.)) (Fig: 334) suas, 5 a -)ai-ee a Se Semidalis

SUBFAMILY ALEUROPTERYGINAE
Aleuropteryx Low, 1885

Scape and pedicel elongate. Male pedicel with a distinct ventral spine. Forewings short,
subtriangular, often rather dark in color. In forewing vein CuP generally sinuous (Fig. 331). In hind
wing veins M and Cu, in their proximal half very close to each other. It is still highly probable that
new species are to be found, and identification usually requires investigation of the male genitalia. In
addition, sclerotized parts of female genitalia provide good specific characters.

This is a Holarctic genus with 16 species recorded from North and Central America, with a center
of distribution in the southwestern U.S.A. and Mexico. Individuals are only very rarely encountered in
great numbers.

KEY TO COSTA RICAN MALE ALEUROPTERYX

1’. Scape about twice as long as broad; spine on pedicel shorter than pedicel annulus ...............4 4. rugosa

Aleuropteryx clavicornis Meinander, 1995
(Figs. 329, 335-338)

DIAGNOSIS. — The male is readily recognized by the long scape and the very prominent spine of
the pedicel. Internal male genitalia: lateral view, Fig. 335; ventral view, Fig. 336; caudal view, Fig.
337; dorsal view, Fig. 338.

GEOGRAPHICAL DISTRIBUTION. — The only known specimen is from Costa Rica:
GUANACASTE.

ALTITUDINAL DISTRIBUTION. — Unknown.

ADULT FLIGHT PERIOD. — March.

HABITAT PREFERENCE. — Unknown.

229 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES
Volume 53, No. 12

Aleuropteryx rugosa Meinander, 1995
(Figs. 339-342)

DIAGNOSIS. — Spiny roof of ring of ninth sternite (IX) of the male genitalia as well as shape of
the process of ninth sternite (pro IX) is diagnostic. Internal male genitalia: lateral view, Fig. 339; ven-
tral view, Fig. 340; caudal view, Fig. 341; dorsal view, Fig. 342.

GEOGRAPHICAL DISTRIBUTION. — This species is known from Nicaragua, and not yet recorded
from Costa Rica.

ALTITUDINAL DISTRIBUTION. — Unknown.

ADULT FLIGHT PERIOD. — In Nicaragua collected from February to September.

HABITAT PREFERENCE. — Unknown.

Neoconis Enderlein, 1929

The head capsule has a large unsclerotized area on the frons, which includes the antennal sockets
and extends downwards medially in a long tongue to the clypeus. In the forewing, the distal crossveins
Sc-R, R,-R,,, and the crossvein-like basal part of R,,; almost forming a line. The radial crossvein
(r-r) of the hind wing strikes the Rs on its branch R, , , well distally of fork of Rs. Identification of spe-
cies requires investigation of internal male genitalia.

This is an American genus with 16 described species, of which two are recorded from Central
America southeast of Mexico.

KEY TO CENTRAL AMERICAN NEOCONIS

1. Forewing almost concolorous; ninth segment of male abdomen apically with about five pairs of very broad dentiform
SOUNDS (HGS; SASSY MD Wee ita alee ee cee Sect te eee A CMe Sete REN Ore Rear ae eee. a GeeereT te N. dentata

1’. Forewing along margin between the veins with round dark spots and round spot around point where crossveins Rs-M
and M-Cu, strike M; ninth segment of male abdomen apico-laterally with short, narrow, apophysis which is bent in-
wards\and bears two or three prominent spines (Figs. 349, 350));. 5. ....55..52.....:: N. marginata

Neoconis dentata Meinander, 1972
(Figs. 343-348)

DIAGNOSIS. — Ectoprocts (Fig. 343: ect) of male with about five strong bristles. Male genitalia,
lateral view, Fig. 343; caudal view, Fig. 344; ventral view, Fig. 345. Paramere, lateral view, Fig. 346.
Internal genitalia ventral view, Fig. 347. Wings, Fig. 348.

GEOGRAPHICAL DISTRIBUTION. — Recorded from Costa Rica (Bataan); and Nicaragua.

ALTITUDINAL DISTRIBUTION. — Unknown.

ADULT FLIGHT PERIOD. — Two records, one from April, the other from July.

HABITAT PREFERENCE. — Unknown.

Neoconis marginata Meinander, 1972
(Figs. 349-355)

DIAGNOsIS. — The spotted forewings (Fig. 355) are distinct, but study of the male genitalia is al-
ways needed to confirm identification. Male genitalia ventral view, Fig. 349; lateral view, Fig. 350.
Penis dorsal view, Fig. 351; lateral view, Fig. 352. Internal male genitalia dorsal view, Fig. 353.
Paramere and stylus lateral view, Fig. 354.

GEOGRAPHICAL DISTRIBUTION. — Recorded from southwestern U.S.A., Mexico, and Nicara-
gua.

230 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES
Volume 53, No. 12

ALTITUDINAL DISTRIBUTION. — Unknown.

ADULT FLIGHT PERIOD. — The records from Nicaragua are from February to September; from
Texas N. marginata is recorded from April to November.

HABITAT PREFERENCE. — Unknown.

SUBFAMILY CONIOPTERYGINAE
Coniopteryx Curtis, 1834

By far the largest genus of dustywings, Coniopteryx has a worldwide distribution and is repre-
sented on all larger land masses. The genus is divided into six subgenera with more than 150 currently
recognized species. This genus is recognizable by the unforked media (Fig. 333M) of the hind wing.
The wing shape is also distinct, with forewings being broad, about twice as long as broad, fore margin
proximally concave, and hind wings with distinctly rounded hind margin. There is a distinct sexual di-
morphism in the antennae. The flagellar segments of males are wider than long and have scalelike
hairs at the apex, while segments of females are longer than wide and only ordinary hairs are present.
The male antennae thus have a much thicker appearance.

KEY TO MALE CONIOPTERYX

1. | Gonarcus (gc) and hypandrium (hy) synscleritous (Fig. 361)... . . (Scotoconiopteryx) = - sea 3
I) Gonarcus/andihypandrium) disclerntous (Fig. 356)! ] 4-5 yf 2) a) ee ys ee 2
2(1'). Base of stylus (sty) attached to gonarcus (gc) far basally of apex (Fig. 357). . (Xeroconiopteryx) .. C. (X.) falcata
Zi Base of stylus attached to gonarcus near apex (Fig. 398)... ... (Contopteryx (Ss: Si7s)) eee ee 8

3(1). The median apical incision of hypandrium (mai) small, less than half the length of hypandrium (Fig. 367) C. (S.) flinti

3’. The median apical incision of hypandrium (mai) at least half as deep as length of hypandrium (Fig. 379)... .. . 4
4(3'). Hypandrium in ventral view with very long and narrow processes terminales (pt) (Fig. 379) . . . . C. (S.) isthmicola
4’. Processes terminales of hypandrium (pt) in ventral view short and blunt (Fig. 384) ................. 5
5(4'). Parameres (pa) without long apical downward-directed part (Fig.381) .............. C. (S.) panamensis
5’. Parameres apically curved downwards, the downward-directed part being about half as long as the horizontal part
(HigSS ORV PON Sacre ay eee 2 LEMS i ee ee ee 6
6(5’). Downwards-directed apical part of paramere strongly curved .................... C. (S.) fumicolor
6) Downwards-directed/apicall parviok paramere straight(Pies 362) 5 2 aes. ae 2) ee eee ee 7
7(6'). Median apical incision of hypandrium (mai) about as broad as deep (Fig. 364) ........... C. (S.) cyphodera
7’. Median apical incision of hypandrium several times as deep as broad (Fig. 374) ............ C. (S.) fumata
SC) a Nobooksonmironsmaxdllanys pal pim Ov brOaG CDEC mien mens cee nnn nee C. (S.) freytagorum
8’. Frons with a distinct hook; fourth segment of maxillary palpi broadened (Fig. 386) ................. 9

9(8'). Hypandrium with an inner, transverse distally bifurcate plate (hyt) proximal to median apical incision (Figs. 387,

388); paramere apically terminating in a fairly long downwards-directed part (Fig. 389)...... C. (C.) californica
9’. Hypandrium without an inner transverse plate (Fig. 398); paramere apically terminating in dorsal transverse plate
(SN SIs) eee oe eee ee ee MOR me aire eon Siseo SB oS C. (C.) mexicana

Coniopteryx (Xeroconiopteryx) Meinander, 1972

This subgenus has a worldwide distribution and species are often confined to low vegetation in
arid areas. The wing membrane is generally pale. In the male genitalia the gonarcus (gc) and

PENNY: LACEWINGS OF COSTA RICA 231

hypandrium (hy) are distinctly discleritous and the styli (sty) are attached to the base of the gonarcus.
Styli are mostly bifurcate with the anterior branch forming an arch below the parameres. Five species
are presently recognized from the Americas, four of which are known from North America.

Coniopteryx (Xeroconiopteryx) falcata Meinander, 1995
(Figs. 356-360)

DIAGNOSIS. — This is the only American species of Xeroconiopteryx, which has scalelike hairs
on the apex of the flagellar segments. A secure identification requires study of the internal male geni-
talia. Male genitalia, lateral view, Fig. 356; caudal view, Fig. 357. Hypandrium, ventral view, Fig.
358. Gonarcus, styli and internal genitalia, ventral view, Fig. 359. Paramere, Fig. 360.

GEOGRAPHICAL DISTRIBUTION. — Described from Nicaragua.

ALTITUDINAL DISTRIBUTION. — Unknown.

ADULT FLIGHT PERIOD. — January and February.

HABITAT PREFERENCE. — Unknown.

Coniopteryx (Scotoconiopteryx) Meinander, 1972

An American subgenus with 28 presently recognized species, five of which are recorded from
Central America. The species are mostly confined to the canopies of trees. Membrane of wings is gen-
erally darkish. In the male genitalia, the gonarcus and hypandrium are proximally synscleritous and
the styli, which form a simple semicircular band below the parameres, are attached to the apex of the
gonarcus.

Coniopteryx (Scotoconiopteryx) cyphodera Johnson, 1978
(Figs. 361-365)

DIAGNOSIS. — Penis (p) very thin and straight. Resembles C. fumata closely, from which it can
be distinguished by the characters mentioned in the key. Male genitalia, lateral view, Fig. 361; caudal
view, Fig. 362. Hypandrium, ventral view, Fig. 364. Internal genitalia, lateral view, Fig. 363; ventral
view, Fig. 365.

GEOGRAPHICAL DISTRIBUTION. — Nicaragua and Panama.

ALTITUDINAL DISTRIBUTION. — Unknown.

ADULT FLIGHT PERIOD. — February to October.

HABITAT PREFERENCE. — Unknown.

Coniopteryx (Scotoconiopteryx) flinti Meinander, 1975
(Figs. 366-370)

DIAGNOSIS. — Sclerotized parts of penis in lateral view relatively thick. Close to C. panamensis,
from which it can be distinguished by the shallow median apical incision of hypandrium (mai) and in
ventral view the distinct processes terminales of hypandrium (pt). Male genitalia in lateral view, Fig.
366; caudal view, Fig. 367. Hypandrium in ventral view, Fig. 369. Internal genitalia in lateral view,
Fig. 368; ventral view, Fig. 370.

GEOGRAPHICAL DISTRIBUTION. — Mexico, Nicaragua, Colombia, and Brazil.

ALTITUDINAL DISTRIBUTION. — Unknown.

ADULT FLIGHT PERIOD. — May.

HABITAT PREFERENCE. — Unknown.

232 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES
Volume 53, No. 12

Coniopteryx (Scotoconiopteryx) fumata Enderlein, 1907
(Figs, 3712375)

DIAGNOSIS. — Penis very thin and straight. Closely resembles C. cyphodera, from which it can
be distinguished by the characters mentioned in the key. Male genitalia in lateral view, Fig. 371; in
caudal view, 372. Hypandrium, ventral view, Fig. 374. Internal genitalia, lateral view, Fig. 373; ven-
tral view, Fig. 375.

GEOGRAPHICAL DISTRIBUTION. — Costa Rica, Dominican Republic, Colombia and Brazil.
Within Costa Rica collected at CARTAGO: Turrialba (C.A.T.LE.).

ALTITUDINAL DISTRIBUTION. — Turrialba is at mid-elevation (1000—1500 m).

ADULT FLIGHT PERIOD. — From Costa Rica recorded in June.

HABITAT PREFERENCE. — Unknown.

Coniopteryx (Scotoconiopteryx) fumicolor Meinander, 1972

DIAGNOSIS. — Part of a group of species with parameres apically downward-directed. The
paramere apices of C. (S.) fumicolor are stronly recurved. The medial notch of the hypandrium is
deeply and narrowly parallel-sided (see Meinander 1972, fig. 148).

GEOGRAPHICAL DISTRIBUTION. — Known only from Costa Rica. Within the country reported
from: PUNTARENAS, Palmar; and SAN JOSE, Perez Zeledon.

ALTITUDINAL DISTRIBUTION. — Unknown.

ADULT FLIGHT PERIOD. — August.

HABITAT PREFERENCE. — The Palmar specimens are labelled as “rain forest.”

Coniopteryx (Scotoconiopteryx) isthmicola Meinander, 1972
(Figs. 376-380)

DIAGNOSIS. — Readily recognized by the very long processes terminales of the hypandrium (pt).
Male genitalia lateral view, Fig. 376; caudal view, Fig. 377. Hypandrium, ventral view, Fig. 379.
Paramere, lateral view, Fig. 378. Parameres, styli and penis sclerites, ventral view, Fig. 380.

GEOGRAPHICAL DISTRIBUTION. — Recorded from Mexico, Honduras, Nicaragua, and Panama.

ALTITUDINAL DISTRIBUTION. — Unknown.

ADULT FLIGHT PERIOD. — March to December.

HABITAT PREFERENCE. — Unknown.

Coniopteryx (Scotoconiopteryx) panamensis Meinander, 1974
(Figs. 381-385)

DIAGNOSIS. — Sclerotized parts of penis in lateral view relatively thick and slightly curved.
Close to C. flinti, from which it can be distinguished by the very broad, median apical incision of the
hypandrium (mai) and the indistinct processes terminales of the hypandrium. Male terminalia, lateral
view, Fig. 381; caudal view, Fig. 382. Hypandrium, ventral view, Fig. 384. Internal genitalia, lateral
view, Fig. 383; ventral view, Fig. 385.

GEOGRAPHICAL DISTRIBUTION. — Costa Rica, Panama, Colombia, Venezuela, Trinidad and
Brazil.

ALTITUDINAL DISTRIBUTION. — Unknown.

ADULT FLIGHT PERIOD. — In Central America recorded in February, March, and May.

HABITAT PREFERENCE. — Unknown.

PENNY: LACEWINGS OF COSTA RICA 233,

Coniopteryx (Coniopteryx) sensu strictu Curtis, 1834

This subgenus has a worldwide distribution with 22 species recorded from America, three of
which are from Central America, southeast of Mexico. Wing membrane is generally pale. In the male
genitalia the gonarcus and hypandrium are distinctly discleritous and the styli are attached to the apex
of the gonarcus (or nearly so). Styli are forked (Fig. 389), the anterior branch of which is attached to
the processus ventralis of the parameres. Hypandrium in lateral view about as high as broad. Processes
laterales of the hypandrium are prominent, forming the dorso-caudal corner of the hypandrium.

Coniopteryx (Coniopteryx) californica Meinander, 1974
(Figs. 386-391)

DIAGNOSIS. — This species belongs to a species group recognized by a hook on the frons, above
the hook a transverse band of stiff hairs and by a broadened fourth segment of the maxillary palpi
(head, Fig. 386). In C. californica, there are no apophyses on any segment of the maxillary palpi. The
parameres terminate apically ina ventral, long, acute apophysis. Male genitalia, lateral view, Fig. 387;
caudal view, Fig. 388. Hypandrium, ventral view, Fig. 390. Internal genitalia, lateral view, Fig. 389;
ventral view, Fig. 391.

GEOGRAPHICAL DISTRIBUTION. — U.S.A. (California and Texas) and Nicaragua.

ALTITUDINAL DISTRIBUTION. — Unknown.

ADULT FLIGHT PERIOD. — In Nicaragua collected in September.

HABITAT PREFERENCE. — Unknown.

Coniopteryx (Coniopteryx) freytagorum Johnson, 1978
(Figs. 392-396)

DIAGNOsIS. — No hook on frons, palpi normal, transverse plate of hypandrium small. An inves-
tigation of the male genitalia is necessary for an accurate identification, although this is presently the
only species of the subgenus Coniopteryx recorded from the isthmus of Central America that lacks a
hook on the frons. Male genitalia, lateral view, Fig. 392; caudal view, Fig. 393. Hypandrium, ventral
view, Fig. 395. Internal genitalia, lateral view, Fig. 394; ventral view, Fig. 396.

GEOGRAPHICAL DISTRIBUTION. — A common species in Honduras, Nicaragua and Costa Rica.
Within Costa Rica collected from GUANACASTE: 3 km SE Naranjo; PUNTARENAS : La Amistad
National Park; SAN JOSE: Escazt.

ALTITUDINAL DISTRIBUTION. — 1500-2330 m (the latter at La Amistad National Park).

ADULT FLIGHT PERIOD. — Throughout the year.

HABITAT PREFERENCE. — Collected on understory saplings, including many Peperonia plants,
in mixed montane primary forest with many large Ficus trees.

REMARKS. — Earlier reports of Coniopteryx simplicior from Costa Rica actually pertain to this
species (see Meinander 1995).

Coniopteryx (Coniopteryx) mexicana Meinander, 1974
(Figs. 397-401)

DIAGNOSIS. — This species belongs to a species group recognized by a hook on the frons; above
the hook a transverse band of stiff hairs; and by a broadened fourth segment of the maxillary palpi. In
C. mexicana there are no apophyses on any segment of the maxillary palpi. Parameres terminate api-
cally in dorsal transverse plates. Male genitalia, lateral view, Fig. 397; caudal view, Fig. 398.

PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES
Volume 53, No. 12

No
ies)
&

Hypandrium, ventral view, Fig. 400. Paramere, lateral view, Fig. 399. Internal genitalia, ventral view,
Fig. 401.
GEOGRAPHICAL DISTRIBUTION. — Mexico, Nicaragua, Costa Rica, and Colombia.
ALTITUDINAL DISTRIBUTION. — Unknown.
ADULT FLIGHT PERIOD. — In Central America this species is recorded from November to May.
HABITAT PREFERENCE. — Unknown.

Semidalis Enderlein, 1905

A genus distributed worldwide with representatives on all larger land masses. Currently about 60
species are recognized, about half of which are American. The species can be distinguished by the in-
ternal male genitalia. There is a certain variation in the genitalia and probably some of the presently
recognized species will prove conspecific when more material is available and the range of variation
better known.

The genus can be distinguished by the forked media (M) in the hind wing and crossvein M-CuA
striking the posterior branch of M and obliquely oriented relative to the longitudinal veins. The shape
of the wings is distinctive, forewings triangular with almost straight fore margin and hind margin to
the tip of CuA. Hind margin of hind wing convex.

KEY TO COSTA RICAN MALE SEMIDALIS

1. Paramere (Fig. 403) dorsally with a forward-directed tooth; uncini fused into an H-shaped transverse plate in caudal

WHEW (TOA A eee oat co uhces lor wapaten cy uci Beads ere Sule, o (oy Geigy tae Oe eee S. arnaudi
1’. Paramere without a dorsal, forward-directed tooth; uncini not fused toasingle plate. ................ 2
2(1’). Ectoproct terminates in a spine, which has a lobe or tooth on its inner edge (Figs. 411,412). ............ 3
2’. Ectoproct blunt and without any appendix on its inner edge (Figs. 418,422) ..................... 4
3(2)sEctoprocticaudallyderminatino invanjacuteispme\(Fie5430)) > 2. 2 2 2 ae eee S. soleri
3’. Ectoproct on its inner edge with an inwards directed blunt lobe (Fig. 424) .............. S. rondoniensis
3”. Ectoproct with a small tooth about medially on its inner margin (Fig. 408) .............. S. manausensis

4(2'). Paramere caudally curved upwards and terminating in a narrow, tapering transverse plate (Fig. 420) . S. panamensis
4’. Paramere caudally straight or tip slightly curved downwards (Fig.415).................. S. mexicana

Semidalis arnaudi Meinander, 1972
(Figs. 402-406)

DIAGNOSIS. — The forward-directed tooth of the paramere (Fig. 403) is distinct. Parameres, in
lateral view, apically swollen, terminally with an upwards directed spine. Uncini (Figs. 404405)
fused into a single transverse plate, which in caudal view has a deep, centro-dorsal incision and a shal-
lower ventral one. Male genitalia, lateral view, Fig. 402. Parameres, lateral view, Fig. 403; ventral
view, Fig. 406. Uncini, caudal view, Fig. 404; lateral view, Fig. 405.

GEOGRAPHICAL DISTRIBUTION. — U.S.A. (Arizona), Mexico and Nicaragua.

ALTITUDINAL DISTRIBUTION. — Unknown.

ADULT FLIGHT PERIOD. — The Nicaraguan specimens were collected from September to
March.

HABITAT PREFERENCE. — Unknown.

235 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES
Volume 53, No. 12

Semidalis manausensis Meinander, 1980
(Figs. 407-413)

DIAGNOsIS. — There are at least three pairs of sclerotized rods below the apex of the parameres,
which apically are forked into two or three spines. Male genitalia, lateral view, Fig. 407; caudal view,
Fig. 408. Caudal parts of internal genitalia, lateral view, different specimens, Figs. 409, 411-413;
ventral view, Fig. 410.

GEOGRAPHICAL DISTRIBUTION. — Mexico south to Brazil (Amazonas) and Peru (Hudanaco).
This species seems to be common in Costa Rica, where it is recorded from CARTAGO: Embalse El
Llano; GUANACASTE: La Taboga Forest Reserve, 9 km SW Cafias; LIMON: Hitoy-Cerere Bio-
logical Reserve; PUNTARENAS: La Amistad National Park; Fila Cedro; Rio Coton; and SAN JOSE:
Escazu.

ALTITUDINAL DISTRIBUTION. — In La Amistad National Park, near Las Alturas, this species has
been found between 1350 and 2100 m.

ADULT FLIGHT PERIOD. — Recorded in Costa Rica from September to March.

HABITAT PREFERENCE. — Individuals were collected in primary forest by beating the understory
vegetation near the edge of La Amistad National Park. This is an area in the rain shadow of the
Talamanca Mountains and in February receives very little rainfall. The forest, although evergreen, is
very dry during this period.

Semidalis mexicana Meinander, 1972
(Figs. 414-419)

DIAGNOSIS. — The ectoprocts are short and blunt. Sclerites below apex of paramere are small.
Male genitalia, lateral view, Fig. 414; caudal view, Fig. 415. Internal genitalia, lateral view, different
specimens, Figs. 416-419.

GEOGRAPHICAL DISTRIBUTION. — Semidalis mexicana has been recorded under different names
from Mexico, Nicaragua, Costa Rica, and Colombia. Within Costa Rica, this species is recorded from
GUANACASTE and SAN JOSE Provinces.

ALTITUDINAL DISTRIBUTION. — Unknown.

ADULT FLIGHT PERIOD. — Within Costa Rica this species is recorded from February to Septem-
ber.

HABITAT PREFERENCE. — Unknown.

Semidalis panamensis Meinander, 1974
(Figs. 420-422)

DIAGNOSIS. — Resembles S. rondoniensis from which it can be distinguished by the lack of the
lobe on the inner margin of the ectoprocts. Except for a pair of lateral sclerites around the apex of the
parameres, the paramere apparently has two backward-directed spines ventrally on the apex. They
can also be separate sclerites. Parameres are caudally curved upwards and terminate in longitudinal,
upward-tapering transverse plates. Semidalis brasiliensis Meinander, 1974 and S. ecuadoriana
Meinander, 1983 are probably conspecific with this species. Male genitalia, caudal view, Fig. 420;
lateral view, Fig. 421. Parameres, lateral view, Fig. 422.

GEOGRAPHICAL DISTRIBUTION. — Costa Rica, Panama, Colombia, Venezuela, Trinidad, and
Brazil (Rio de Janeiro). If S. brasiliensis and S. ecuadoriana prove conspecific, there are records also
from Colombia, Ecuador, and Brazil (Santa Catarina). Within Costa Rica this species was collected at
PUNTARENAS: Quepos.

ALTITUDINAL DISTRIBUTION. — Quepos is at sea level.

236 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES
Volume 53, No. 12

ADULT FLIGHT PERIOD. — The Costa Rican collection was in February.
HABITAT PREFERENCE. — Unknown.

Semidalis rondoniensis Meinander, 1982
(Figs. 423-429)

DIAGNOSIS. — Typical for S. rondoniensis is the lobe on the inner margin of the ectoproct. There
is aconsiderable variation in the genitalia and in some specimens the lobe is longer and the apex is ser-
rate. Male genitalia, lateral view, Fig. 423; caudal view, Fig. 424. Parameres, lateral view, different
specimens, Figs. 425-429.

GEOGRAPHICAL DISTRIBUTION. — Costa Rica, Colombia, Venezuela, Peru, Bolivia and Brazil.
Within Costa Rica, S. rondoniensis is recorded from LIMON: Hitoy-Cerere Biological Reserve.

ALTITUDINAL DISTRIBUTION. — Elevation ca. 200 m.

ADULT FLIGHT PERIOD. — March.

HABITAT PREFERENCE. — Understory of low, moist, open, secondary vegetation along trail.

Semidalis soleri Monserrat, 1985
(Figs. 430-433)

DIAGNOSIS. — The male genitalia resemble those of S. rondoniensis, but the very acute
ectoprocts are distinct. Male genitalia, lateral view, Fig. 430; caudal view, Fig. 431. Internal genitalia,
lateral view, Fig. 432. Caudal parts of internal genitalia, ventral view, Fig. 433.

GEOGRAPHICAL DISTRIBUTION. — Semidalis soleri was described from Mexico. There is a long
series from Costa Rica; PUNTARENAS: La Amistad National Park.

ALTITUDINAL DISTRIBUTION. — Records from near Las Alturas, La Amistad National Park in
Puntarenas Province indicate a elevational distribution from 1500 to 2000 m.

ADULT FLIGHT PERIOD. — February to March.

HABITAT PREFERENCE. — Individuals of this species were collected 1n primary forest by beating
the understory vegetation near the edge of La Amistad National Park. This is an area in the rain
shadow of the Talamanca Mountains and in February receives very little rainfall. The forest, although
evergreen, is very dry during this period.

FAMILY DILARIDAE

by
Norman D. Penny

Dilarids superficially resemble small moths and many go undetected in light trap samples. New
World species all belong to the genus Nallachius, which has broad dark bands or large spots on the
wings. Males have pectinate (feathery) antennae (Fig. 434), whereas females have the more usual
filiform antennae. Larvae are predators of wood-dwelling arthropods in rotting logs. Three species
have been recognized in the Costa Rican fauna, but because of their small size and forest habitat, few
individuals have been collected. They are attracted to light and that is the most frequent means of cap-
ture.

KEY TO COSTA RICAN NALLACHIUS

|. Forewing MP, fused with CuA for some distance; MP, appearing simple to near wing apex. (Fig. 436)... ... . 2
\'. Forewing MP, not fused with CuA; MP, appearing deeply forked(to basad of Rs + MA fork) (Fig. 437) N. pulchellus

PENNY: LACEWINGS OF COSTA RICA 237

2(1). Dark bands of forewing mostly complete (Fig. 436); dorsal lobes of ectoproct pincer-like ......... N. parkeri
2'._ Dark bands of forewing mostly incomplete, giving appearance of nebulous clouds(Fig. 435); dorsal lobes of
EA CHLOcHSiMplesMOUDININCALC. ovacun agi We esac Hen G4 Ss Sale Secs Se A ones eA we N. americanus

Nallachius americanus (McLachlan, 1880)
(Fig. 435)

DIAGNOSIS. — This species can be separated from N. pulchellus by the apparent deeply forked
forewing MP, in the latter species. The only other Costa Rican dilarid, N. parkeri also has a shallowly
forked MP,, and can only be separated with confidence by studying the male genitalia. In general, N.
americanus appears to be a somewhat paler species with slightly narrower wings. However, there is
too much variability to these character states to be consistently useful.

GEOGRAPHICAL DISTRIBUTION. — Eastern U.S.A. south to Venezuela (Adams 1970). Within
Costa Rica, N. americanus has been collected only with light traps at 3 km SE of Rio Naranjo in
GUANACASTE Province.

ALTITUDINAL DISTRIBUTION. — Unknown.

ADULT FLIGHT PERIOD. — Late November to late December.

HABITAT PREFERENCE. — Unknown.

Nallachius parkeri Penny, 1994
(Fig. 436)

DIAGNOSIS. — This species closely resembles NV. americanus, but males can be separated by the
deep bifurcation of the dorsal lobe of the ectoproct and lack of a triangular projection on the posterior
border of the ninth tergite.

GEOGRAPHICAL DISTRIBUTION. — This species is only known from light traps at 3 km SE of Rio
Naranjo in GUANACASTE Province.

ALTITUDINAL DISTRIBUTION. — Unknown.

ADULT FLIGHT PERIOD. — This species may have a bimodal emergence period. Three speci-
mens have been collected from late January to early March, while one male was taken in
mid-September.

HABITAT PREFERENCE. — Unknown.

Nallachius pulchellus Banks, 1938
(Fig. 437)

DIAGNOSIS. — Currently this is the only species of dilarid known from Costa Rica which does
not have MP, fused with CuA, giving MP, a deeply forked appearance.

GEOGRAPHICAL DISTRIBUTION. — The male type is from Cuba. Other specimens are known
from U.S.A. (Arizona) south through Mexico. Costa Rica is as far south as this species has been re-
corded. Within Costa Rica this is the most widely distributed and most commonly encountered spe-
cies of Nallachius. It is known from GUANACASTE: 3 km SE of Rio Naranjo; Santa Rosa National
Park; PUNTARENAS: Reserva Carara; and SAN JOSE: Escazt.

ALTITUDINAL DISTRIBUTION. — Unknown.

ADULT FLIGHT PERIOD. — Early December to May (Rio Naranjo), but a record from farther
north at Santa Rosa National Park in July suggests that adults may be present during most of the year.

HABITAT PREFERENCE. — Unknown.

238 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES
Volume 53, No. 12

FAMILY HEMEROBIIDAE

by
Victor J. Monserrat

Hemerobiids constitute one of the largest families of Neuroptera, with about 550 known species
worldwide, and their geographical distribution includes all continents except Antarctica. This is a rel-
atively abundant family in Costa Rica and is well represented in the fauna. Hemerobiids are me-
dium-sized insects, easily recognized by their filamentous antennae, by wings held roof-like over the
abdomen when resting, and by their generally pale brown, frequently cryptic, coloration. Adults are
weak fliers with crepuscular and noctural habits, and because of this, habitually spend the day resting
on leaves and are frequently attracted to lights at night. They are usually predators of phytophagous in-
sects, and are utilized in biological control of aphids, scale insects, etc., but also occasionally feed on
sugary fluids and pollen. Larvae are also very active predators with broad bodies and curved, impos-
ing mandibles used to capture and siphon body fluids from soft-bodied, phytophagous insects. Larval
development requires three instars and about 15 to 20 days for completion, at which time a small silk
capsule is constructed in which it pupates. Usually species pass through only one or two generations
per year, and occasionally different species are restricted to particular vegetational substrates or alti-
tudes.

KEY TO GENERA OF COSTA RICAN HEMEROBIIDAE

il. Forewing with two radialisectors? (Figs: 5075583))- 2. 2 9. . = 2 1 2) te oe ere een 2D,
i Rorewing: with three‘or more’ iradiallsectorss, (Figs, 4457492)) 2 202 2 ake se 2 3
2) se korewineiwithitworenadate:senes\ (194559) eisi4 ope) oeene el. Gin ee een ete eee een Sympherobius
2), ‘Forewing with only one pradate’'senes (Fic. 507) .- - 2...) 2: se as ae 2 ee Eee 6
3(1). Forewing with two basal crossvems m’subcostal space(Fis. 492) 2 >. 5.2. -5. 4. aes eee Megalomus
3’. Forewing with only one basal crossvein in subcostal space (Figs. 445,530) ...............-+---+-- 4
4(3'). Forewing with two crossveins between last “radial sector” and the radius (Fig. 445) ........... Hemerobius
4’. Forewing with only one crossvein between last “radial sector” and the radius (Fig. 530). .............. 5)
5(4’). Forewing with three gradate series; MP fused with CuA (Fig. 530). .................-..- Nusalala
5'. Forewing with two gradate series; MP connected to CuA by a crossvein (Fig. 504) ............ Micromus
6(2'). First fork of first “radial sector” more distal than first fork of second “radial sector”? ............. Biramis
6’. First fork of first “radial sector” at same level as first fork of second “radial sector”... ........ Notiobiella

Biramis Oswald, 1993

This genus was described in 1993 for a primitive species of hemerobiid from Venezuela. There is
a second, as yet undescribed, species known from Costa Rica.

Hemerobius Linnaeus, 1758

This genus is very speciose, with 144 species worldwide. It is the most widely distributed
hemerobiid genus, being found on all continents except Antarctica, and many islands of the Indian,
Atlantic and Pacific Oceans. Thirty species are known from Central and South America, with 11 spe-
cies found in Costa Rica.

PENNY: LACEWINGS OF COSTA RICA 239

KEY TO COSTA RICAN HEMEROBIUS

1. Ronrewincuwithicubitalcelliopen|(Rigs: 4405446) - 3.22% 3. ccs ey a oe ieee Pee 3
Me Bormowineawitnicubitalcellliclosedl(Riss 44443) 0 es ise ei oe eee eek each cue tl -peceioeii) cursuiceto cod ouge 2

2(1'). Male ectoproct with large ventral lobe (Fig. 457); entoprocesses widely separated at base and more than six times

as long as wide (Fig. 459); with posterodistal margin nearly straight (Fig. 443) ............ H. discretus
ae Male ectoproct with very small ventral lobe (Fig. 474); entoprocesses close at base and only four times as long as
wide (Fig. 476); forewing with posterodistal margin more convex (Fig. 448)... .......... H. martinezae

3(1). Forewing elliptical (width/length ratio ~ 0.47), with at most 12 subcostal veinlets in the costal space before the

PLELOS OMAN (EIORA SINE ear ee Mc fee ery eee Si ee Sees ee eae 4
Bh Forewing more elongate (width/length ratio ~ 0.42), with more than 12 subcostal veinlets in the costal space before
TOES IIA IO SRAAO) bio cs ays lea eas SPM a sok OS Rae eben Gebel Cokero eel REA? Se EROURR Nene 7
4(3). Forewing with a dark brown spot over base of costal space (Fig.451) ................... H. pennii
4’. Forewing without dark brown spot over base of costal space (Fig. 452) ...............---+..-.-.- 5

5(4'). Scape yellow, with a brown stripe on the external side; thorax dark brown without medial yellow band ..... .
a are eae ae Bs Sle hiclihn Rael hie ens oe A ee ae H. withycombei

ie Scape yellow, without brown stripe on external surface; thorax dark brown with a medial yellow band .... . 6

6(5'). Basal flagellomeres of antenna flattened (Fig. 439) in male; wings elliptical (width/length ratio = 0.45) (Fig. 450)
Ry ere rts Nos sete ee Ee ei ayam, = ate: See Mgsegs Ge CERES wal & BER ION H. hirsuticornis

7(3'). Thorax brown without a pale or yellow band over the central medial zone; pterostigma distinct (Fig. 449)... . .
50 0 6 6180 ONCE Rn Seas eR CG at ee eo CT emer tS ce Od ore omc H. nigridorsus

1h Thorax brown with a pale or yellow band over the central medial zone; pterostigma inconspicuous ...... . 8
Se aeborewine withiinner eradate veins aligned (Fig: 447) 2 ee ee H. jucundus
8’. Forewing with inner gradate veins not aligned (Fig. 446). ........-...2-+:++++++5-++-.----- 9

9(8'). Forewing with the costal space strongly dilated basally (three times as wide basally as apically) (Fig. 446)... . .
sa 5 ae ote oS ee rr near Warn orton Gate tartan okra coe H. hernandezi

uy Forewing with costal space somewhat dilated basally (twice as wide basally as apically) (Fig. 445) ....... 10
10(9). Head brown, with darker brown spots on the vertex, frons and external surface of scape .......... H. gaitoi
10’. Head brown, without darker brown spots on the vertex, frons and external surface of scape ....... H. bolivari

Hemerobius bolivari Banks, 1910
(Figs. 442, 453-456)

DIAGNOSIS. — This is one of the most variable species in the genus, both in size and pigmenta-
tion, in venation, and genitalia. The wings are lanceolate and basically brown colored, especially the
forewings, with slightly darker markings along the gradates and basally. The male genitalia (Fig.
453456) can help with differentiation.

GEOGRAPHICAL DISTRIBUTION. — A very broadly distributed species from Mexico, Guatemala,
Costa Rica, Panama, Colombia, Venezuela, Ecuador, Peru, Bolivia, Paraguay, Brazil, Uruguay,
Chile, and Argentina. In Costa Rica this species is known from: GUANACASTE, Canas; Rio
Naranjo; PUNTARENAS, Monteverde; Bellavista; La Amistad National Park; Las Alturas; and SAN
JOSE, Escazi.

ALTITUDINAL DISTRIBUTION. — Sea level to 3100 m, in Costa Rica to 1500 m.

240 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES
Volume 53, No. 12

ADULT FLIGHT PERIOD. — This species has been collected in all months of the year; Costa Rican
records are from January to June, September, and December.

HABITAT PREFERENCE. — This species is known from a multitude of habitats, generally dry,
sunny, open areas and frequently associated with human activity. They are usually found on trees.

Hemerobius discretus Navas, 1917
(Figs. 443, 457-459)

DIAGNOSIS. — Large winged (forewing length 9.2—13.5 mm); this is one of the largest brown
lacewings in Costa Rica. The forewings are highly variegated with brown, slightly lanceolate and with
a straight posterodistal margin (Fig. 443). Females appear to be somewhat larger. The male external
genitalia are unmistakable (Fig. 457-459).

GEOGRAPHICAL DISTRIBUTION. — U.S.A. (western), Mexico, Guatemala, Costa Rica, and Pan-
ama. In Costa Rica specimens have been encountered at: CARTAGO: Cartago; Chirrip6; Parque
Nacional Irazu; HEREDIA: Parque Nacional Braulio-Carrillo; SAN JOSE, San Gerardo de Dota, San
Jose.

ALTITUDINAL DISTRIBUTION. — Mid-elevations from ca. 1200 to 3000 m.

ADULT FLIGHT PERIOD. — February to December.

HABITAT PREFERENCE. — Generally associated with dry and semiarid zones. Specimens have
been collected on pines and with conifer pollen in their digestive tracts.

Hemerobius edui Monserrat, 1991
(Figs. 444, 460-462)

DIAGNOSIS. — A small species with very rounded forewings, sparse, open venation and varie-
gated brown markings (Fig. 444). It is easily differentiated by the male genitalia (Fig. 460-462).

GEOGRAPHICAL DISTRIBUTION. — This species is known from Mexico, Costa Rica, Colombia,
and Peru. In Costa Rica collections are from: ALAJUELA: (without precise locality);
GUANACASTE, Maritza Biological Station, southwest side of Orosi Volcano; Pitilla Biological Sta-
tion, 7 km S Santa Cecilia; and PUNTARENAS, Monteverde.

ALTITUDINAL DISTRIBUTION. — 600-1500 m (Monteverde).

ADULT FLIGHT PERIOD. — July to September.

HABITAT PREFERENCE. — Subtropical moist forest.

Hemerobius gaitoi Monserrat, 1996
(Figs. 445, 463-466)

DIAGNOSIS. — Somewhat variable in wing and integumentary coloration. It is easily recogniz-
able by the pallid color of the forewings upon which are arranged two dark spots in the basal region
(Fig. 445). The male gentialia (Figs. 463-466) can also serve to differentiate this species.

GEOGRAPHICAL DISTRIBUTION. — Widely distributed in the Neotropical Region: Mexico, Gua-
temala, Costa Rica, Dominican Republic, Venezuela, Ecuador, and Brazil. In Costa Rica it has been
collected at: ALAJUELA (no further locality); PUNTARENAS, La Amistad National Park; Rio
Coton; Rio Bellavista.

ALTITUDINAL DISTRIBUTION. — 1000-2100 m.

ADULT FLIGHT PERIOD. — February and April.

HABITAT PREFERENCE. — Associated with trees in mountainous zones.

PENNY: LACEWINGS OF COSTA RICA 241

Hemerobius hernandezi Monserrat, 1996
(Figs. 446, 467-469)

DIAGNOSIS. — Easily recognized by its broad, oval, dark forewing, which is broadly expanded in
the basal region of the costal space (Fig. 446). The male genitalia (Figs. 467-469) will also serve to
identify this species.

GEOGRAPHICAL DISTRIBUTION. — Broadly distributed from Mexico, Guatemala, Nicaragua,
Costa Rica, Panama, Colombia, Venezuela, Brazil, and Paraguay. In Costa Rica it has been frequently
collected at: ALAJUELA, San Ramon Forest Reserve, Rio San Lorencito; CARTAGO, Embalse El
Llano; GUANACASTE, Rio Naranjo; LIMON: PUNTARENAS, Monteverde; Irazi Vulcano; La
Amistad National Park, Las Alturas; San Vito; and SAN JOSE, Est. Carara.

ALTITUDINAL DISTRIBUTION. — 1000-2200 m.

ADULT FLIGHT PERIOD. — In Costa Rica there appears to be a bimodal emergence pattern from
January to June, and September to December.

HABITAT PREFERENCE. — Humid, forested zones.

Hemerobius hirsuticornis Monserrat, 1999
(Figs. 439, 450, 481-484)

DIAGNOSIS. — This small species has elliptical wings with variegated membrane (Fig. 450) and
the males possess progressively swollen antennal segments in the basal third, forming a unique struc-
ture for this family (Fig. 439). Male genitalia with ectoprocts wide and bearing a preapical denticle
(Figs. 481-484).

GEOGRAPHICAL DISTRIBUTION. — Described from Ecuador, it has been found in Mexico, Hon-
duras, and Costa Rica. Within Costa Rica it is known only from PUNTARENAS: Monteverde.

ALTITUDINAL DISTRIBUTION. — Elevation ca. 1500 m.

ADULT FLIGHT PERIOD. — September.

HABITAT PREFERENCE. — The specimen from Monteverde was collected along the trails below
the Pension Quetzal. This is a low, secondary, scrub forest along the margin of pastures. It is relatively
open and subject to winds and rapidly changing temperatures.

Hemerobius jucundus Navas, 1927
(Figs. 447, 470-473)

DIAGNOSIS. — Easily recognizable by the gradate veins of the forewing, which are arranged in
regular, aligned, step-wise progression (Fig. 447)—not in a broken line, as in other species. The male
genitalia (Figs. 470-473) can also serve to differentiate this species.

GEOGRAPHICAL DISTRIBUTION. — Known from Mexico, Costa Rica, and Panama. In Costa Rica
it has been collected at: CARTAGO, La Georgina; Villa Mills.

ALTITUDINAL DISTRIBUTION. — 1200-2740 m.

ADULT FLIGHT PERIOD. — March to May, and July to October. Within Costa Rica it has only
been collected in August.

HABITAT PREFERENCE. — Associated with dry and semidesert zones.

Hemerobius martinezae Monserrat, 1996
(Figs. 448, 474-476)

DIAGNOSIS. — Brown head with darker stripes on the vertex and external surface of the antennal
scape. Wings as in Figure 448. The male genitalia are unmistakable (Figs. 474-476).

242 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES
Volume 53, No. 12

GEOGRAPHICAL DISTRIBUTION. — Mexico, Guatemala, and Costa Rica. In Costa Rica it is abun-
dant, collected at: CARTAGO, Irazu Vulcano; HEREDIA, Est. Barva; PUNTARENAS, La Amistad
National Park; Las Alturas; and Las Tablas.

ALTITUDINAL DISTRIBUTION. — High elevations up to 2900 m, most common at about 2000 m.

ADULT FLIGHT PERIOD. — This species can be collected at all times of year, and in Costa Rica
has been taken from December to April, June, and October.

HABITAT PREFERENCE. — Associated with montane forest vegetation, frequently in oak
(Quercus) forests.

Hemerobius nigridorsus Monserrat, 1996
(Figs. 449, 477-480)

DIAGNOSIS. — Easily recognized by the dark brown pterostigma (Fig. 449), and the dorsal re-
gion of the thorax without a medial yellow band, as in the other species. The male genitalia (Figs.
477-480) will also serve to differentiate this species.

GEOGRAPHICAL DISTRIBUTION. — Known only from Costa Rica: PUNTARENAS, La Amistad
National Park, Las Alturas.

ALTITUDINAL DISTRIBUTION. — 1500-1600 m.

ADULT FLIGHT PERIOD. — February and March.

HABITAT PREFERENCE. — Collected on understory saplings, including many Peperonia plants,
in mixed montane primary forest with many large Ficus trees at a dry period of the year in a mountain-
ous rain shadow area.

Hemerobius pennii Monserrat, 1996
(Figs. 451, 485-488)

DIAGNOSIS. — Very small species (5 mm forewing length), characterized by spots on the
forewing which extend to the costal field (Fig. 451). The species is also easily distinguished by the
male genitalia (Figs. 485-488).

GEOGRAPHICAL DISTRIBUTION. — Known from Costa Rica and Ecuador. In Costa Rica it is
known from: ALAJUELA, 20 km S Upala; PUNTARENAS, La Amistad National Park, Las Alturas.

ALTITUDINAL DISTRIBUTION. — 870-1500 m.

ADULT FLIGHT PERIOD. — February to July.

HABITAT PREFERENCE. — Collected from understory saplings, including many Peperonia
plants, in mature montane primary forest with many large Ficus trees at a dry period of the year ina
mountainous rain shadow area.

Hemerobius tibialis Navas, 1917

This species was originally described from Colombia and subsequently reported from Costa
Rica. However, the original description included few distinguishing characteristics and the type (in
the Apolinar Collection in Bogota, Colombia) was destoyed in a fire in 1948. This species must cur-
rently be considered a nomen dubium. For a fuller discussion of this species see Monserrat (1996).

Hemerobius withycombei (Kimmins, 1928)
(Figs. 452, 489-491)

DIAGNOSIS. — Very rounded wings with sparse, open venation and very little pigmentation, ex-
cept at the pterostigma, which is very dark and distinctive (Fig. 452). The body coloration is pale

PENNY: LACEWINGS OF COSTA RICA 43

brown and emphasizes the yellow antennae with lateral brown band on the scape. Male genitalia as in
Figs. 489-491.

GEOGRAPHICAL DISTRIBUTION. — Known from Mexico, Costa Rica, and Colombia. In Costa
Rica this species has been collected at: GUANACASTE, Maritza Biological Station, southwest side
of Orosi Vulcano.

ALTITUDINAL DISTRIBUTION. — 550 m.

ADULT FLIGHT PERIOD. — March to September elsewhere, but only in July in Costa Rica.

HABITAT PREFERENCE. — Mature forests, probably associated with conifers.

Megalomus Rambur, 1842
This genus contains about 40 known species. It is widely distributed in Europe, North Africa, the
Americas and Palearctic Asia, sparsely distributed in Oriental Asia (Taiwan and India) and is absent
from the A frotropical and Australian Regions. Mega/omus 1s broadly distributed in Central and South

America, with two species in Costa Rica.

KEY TO COSTA RICAN MEGALOMUS

1. Hind wing with MA reaching Rs, before its first fork (Fig. 494); forewing ovoid, with 4—5 radial sectors; first

subcostal crossvein very oblique with respect to the second (Fig. 492) .................... M. minor
1’. Hind wing with MA reaching Rs, after its first fork (Fig. 495); forewing apically tapered, even falcate, with 9-11 ra-
dial sectors; first subcostal crossvein parallel to the second (Fig. 493)... ...............04. M. pictus

Megalomus minor Banks, 1905
(Figs. 492, 494, 496-499)

DIAGNOSIS. — A small species with pale integument and pale antennae, and generally with
scape and pedicel darker externally. Forewings elliptical, rounded at its apex, with the first crossvein
of the subcostal field oblique (Fig. 492), membrane pale with variegations and dispersed darker spots.
Male genitalia as in Figures 496-499.

GEOGRAPHICAL DISTRIBUTION. — Widely distributed around the Caribbean, northward to
U.S.A. (AL, CA, MS, NV, see Penny et al. 1997) southward to Colombia. Within Costa Rica it has
been collected in all provinces: ALAJUELA, CARTAGO, GUANACASTE, HEREDIA, LIMON,
PUNTARENAS, and SAN JOSE.

ALTITUDINAL DISTRIBUTION. — Within Costa Rica this species has been collected at 200,
680-700, and 1400 m. It has been collected from sea level to 1400 m in extralimital areas.

ADULT FLIGHT PERIOD. — Adults present year round, but in Costa Rica are most abundant from
February to April.

HABITAT PREFERENCE. — Open fields and grasslands, also in tropical forest zones and coffee
plantations.

Megalomus pictus Hagen, 1861
(Figs. 493, 495, 500-503)

DIAGNOSIS. — A large species (10-12 mm forewing length) with dark brown integument and
brown antennae. Forewings somewhat pointed, in some specimens even markedly falcate, with the
crossveins of the subcostal field parallel (Fig. 493), pale membrane in some specimens, with much
variegation and brown spots in others. Male genitalia as in Figures 500-503.

GEOGRAPHICAL DISTRIBUTION. — Widely distributed in Central America (Mexico, Guatemala,
Honduras, and Costa Rica). In Costa Rica it has been collected at the Irazu Volcano.

244 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES
Volume 53, No. 12

ALTITUDINAL DISTRIBUTION. — 2200 m and 2590-3200 m.

ADULT FLIGHT PERIOD. — February, June to July, and December elsewhere. In Costa Rica
adults are known from December.

HABITAT PREFERENCE. — Probably montane tropical forest. Recorded on Pinus, with conifer-
ous pollen in its digestive tract.

Micromus Rambur, 1842

This is a genus of about 100 described species, with an almost cosmopolitan distribution (Europe,
Asia, Africa, North and Central America, Australia, and many islands in the Atlantic, Pacific and In-
dian oceans). In North America, eight species are recognized, some of which reach Mexico and the
Caribbean. Earlier records of this genus from South America pertain to Nusalala Navas and there ap-
pear to be no confirmed species from that continent. Only one species occurs in Costa Rica.

Micromus subanticus (Walker, 1853)
(Figs. 504—S06)

DIAGNOSIS. — A small, pale brown species, vertex pale with brown spots on both sides and tho-
rax with paler median stripe. Forewing (Fig. 504) elongate, narrow and pale brown, with a few small
brown spots (forewing length 4.5—7.5 mm, hind wing length 4.0—5.5 mm). Male ectoproct and inter-
nal genitalia (Figs. 505, 506). The morphology was described in detail by Klimaszewski and Kevan
(1988).

GEOGRAPHICAL DISTRIBUTION. — A transcontinental species widely distributed in North Amer-
ica (from southern Canada to Mexico), as well as the Dominican Republic, Haiti and Cuba. Previously
unknown from Costa Rica, it is here reported from: ALAJUELA: Upala; GUANACASTE: Canas and
Rio Naranjo; and PUNTARENAS: San Vito.

ALTITUDINAL DISTRIBUTION. — Sea level to about 1200 m.

ADULT FLIGHT PERIOD. — Collected throughout the summer months in temperate climates. In
Costa Rica records are from all months except June, July, and October.

HABITAT PREFERENCE. — Open and mixed forest fields, on birch, vineyards, citrus, alfalfa
fields, but primarily on grass and low vegetation. Information on biology was outlined in
Klimaszewski and Kevan (1988). Usually collected at lights.

Notiobiella Banks, 1909

This genus has a pan-tropical distribution (Central and South America, sub-Saharan Africa,
Southeast Asia, Australia and the Southwestern Pacific) with ca. 35 known species. In the Americas
eight species are known, distributed from Mexico, Central America, and the Caribbean south to
Brazil, Argentina, and Paraguay. Within Costa Rica two species are known.

KEY TO MALES OF COSTA RICAN NOTIOBIELLA

1. Membranous formations of male internal genitalia lacking chitinous denticles (Figs. 508,510) .... . N. rubrostigma
|’. Membranous formations of male internal genital possessing chitinous denticles (Figs. 512-514)... ... N. mexicana

PENNY: LACEWINGS OF COSTA RICA 245

Notiobiella mexicana Banks, 1913
(Figs. 511-514)

DIAGNOSIS. — This is a pale brown species. Wing venation pale brown with forewing longitudi-
nal vein forks and gradate veins darker (Fig. 511). External morphological characters are somewhat
variable and examination of the male genitalia (Figs. 512—514) is needed for reliable identification.
The morphology and genitalia were treated by Monserrat (1984).

GEOGRAPHICAL DISTRIBUTION. — Previously known only from Mexico (San Luis Potosi and
Guadalajara). Reported here from Costa Rica: GUANACASTE: Canas, where it appears quite com-
mon.

ALTITUDINAL DISTRIBUTION. — Sea level to more than 1000 m.

ADULT FLIGHT PERIOD. — Costa Rican specimens have been collected in January, February,
July, and September.

HABITAT PREFERENCE. — It appears to inhabit sunny areas and is frequently collected at lights.

Notiobiella rubrostigma Navas, 1914
(Figs. 507-510)

DIAGNOSIS. — This species is very similar to NV. mexicana in external morphology, pigmentation
and venation (Fig. 507), and the external morphological characters mentioned in the identification
key need to be confirmed with examination of the male genitalia (Figs. 508-510) for confident identi-
fication. The morphology and genitalia were treated by Monserrat (1984).

GEOGRAPHICAL DISTRIBUTION. — Widely distributed in the neotropical region: Mexico, El Sal-
vador, Honduras, Panama, Colombia, Venezuela, Bolivia, Peru, Brazil, Paraguay, and Argentina.
Previously unknown from Costa Rica, but here recorded from GUANACASTE: Rio Naranjo; and
SAN JOSE: La Hermosa.

ALTITUDINAL DISTRIBUTION. — Sea level to 1000 m.

ADULT FLIGHT PERIOD. — Individuals have been collected throughout the year (January to
April, July to September, and November) in other parts of its range; reported in Costa Rica from Janu-
ary, February, April, August, and September.

HABITAT PREFERENCE. — Humid zones to open and sunny areas, usually in arboreal vegetation.

Nusalala Navas, 1913
A Neotropical genus with 17 valid species. It is known from southern Mexico, Central America,
the Caribbean Islands (where it is especially rich in species), and South America (though absent in
most of Amazonia and the southern cone). Replaces Micromus in South America and is a common

tropical element. In Costa Rica Nusalala is represented by seven species.

KEY TO COSTA RICAN NUSALALA

ile Forewing coriaceous and brachypterous, hind wing tiny and scalelike........-..--.--. N. brachyptera
I’. Both fore- and hind wing membranous and extending beyond abdomen .......-. 2-22-25 5-5 2 eee 2
2(1'). Costal space of forewing basally with crossveins present (Fig. 521); male genitalia (Figs. 522-523) . . . N. irrebita

2 ee Castallspace of forewing with crossvems absent (Fig. 527) .- - - 2.2 5 pe ee ee 3

3(2'). Forewing with a dark line through all three gradate series and with a medial dark longitudinal stripe (Fig. 527); male
Emin (Ries, SPARS) Fr oo co alka) wm oltcu ow omomon oat iosd aBino eo SENGioh alias Cuchi ol cienlcl wn lpr suci cua Rcato: hoi N. uncata
3", WWYTuinoyin nese Ayres more Sg eno p06 6 ph 0 on bos comes Olio 6.0.5. buck o e e O a

246 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES
Volume 53, No. 12

4(3'). Forewing: with distinct dark spots (Pigs: 5155530). ==) 5 5. 32 oe 2 2 OR SS See eee 5
A's Forewing without distinct dark spots\(Eigs. S85524)\ 2) Sy 5 ee ee i ee 6
5(4). Forewing subtriangular (Fig. 515), width/length ratio = 0.40; male genitalia (Figs. 516-517) ...... N. championi
5’. Forewing elongated (Fig. 530), width/length ratio = 0.36; male genitalia (Figs. 531-532) ...... N. unguicaudata

6(4’). Forewing with external gradates darker than the other series (Fig. 524). Male genitalia (Figs. 525-526) . N. tessellata
6’. Forewing with mid- and external gradate series darker than inner one (Fig. 518). Male genitalia (Fig. 519-520). . .

soli ete ah gy SE age Sian tau SG Seah cele Sg ay las Ue SUEY syie a ative cy >) ote wees io ae ee N. ghioi
Nusalala brachytpera Oswald, 1996
(Fig. 514a-c)
DIAGNOSIS. — This species can be distinguished from other species of Nusalala by the

coriaceous and brachypterous forewing, the highly reduced and scalelike hind wing, and the
posteroventral spine on the male ectoproct.

GEOGRAPHICAL DISTRIBUTION. — Known only from four localities in Costa Rica: CARTAGO:
Villa Mills, km 97.5 Inter-American Highway, R. K. Colwell, 1 male (Oswald Coll.); km 97 on Cerro
de la Muerte, M. L. Jameson , | male (CAS); 5 km (3 mi) south Empalme, O’Brien, O’Brien, and
Marshall, | female (CAS); SAN JOSE: 46.7 km (29 mi) north of San Isidro del General, C. W. and
LB. O Bren, (male (CAS):

ALTITUDINAL DISTRIBUTION. — 2620-3350 m.

ADULT FLIGHT PERIOD. — May to July.

HABITAT PREFERENCE. — At least some of the specimens have been collected by beating the low
vegetation of the high altitude “paramo.”

REMARKS. — Although this species appears very distinctive, we have seen female specimens of
what may be a second, related species of brachypterous Nusalala.

Nusalala colombiensis (Banks, 1910)

DIAGNOSIS. — This species has only 3 to 4 branches of the “Rs” and the forewing 1s apically lan-
ceolate and pointed with a longitudinal, dark stripe in the middle. These characters will separate this
species from all others known from Costa Rica.

GEOGRAPHICAL DISTRIBUTION. — Originally described from COLOMBIA; here reported from
Costa Rica: SAN JOSE: Zurqui de Moravia, | female (TAMU).

ALTITUDINAL DISTRIBUTION. — 1600 m.

ADULT FLIGHT PERIOD. — September.

HABITAT PREFERENCE. — Unknown.

Nusalala championi Kimmins, 1937
(Figs. 515-517)

DIAGNOsIS. — A large, dark brown spotted species. Fore- and mid-femora and tibiae with two
dark preapical spots. Forewing (Fig. 515) wide, broader at mid-length, and somewhat acute at apex,
hind wing pale brown with darker external gradates, forewing with 5—7 “radial sectors,” dark varie-
gated and irregularly dark dotted, usually with dark dots on medial and outer gradates (forewing
length 9-11 mm, hind wing length 7-9 mm). Male ectoproct and internal genitalia (Figs. 516-517).

GEOGRAPHICAL DISTRIBUTION. — Known only from Mexico, Guatemala, Costa Rica, and Pan-
ama. Within Costa Rica it has been collected in ALAJUELA: San Ramon; CARTAGO: Turrialba;
and PUNTARENAS: Monteverde.

ALTITUDINAL DISTRIBUTION. — 1400 to 1520 m in Costa Rica.

PENNY: LACEWINGS OF COSTA RICA IAT

ADULT FLIGHT PERIOD. — Collected during the first seven months of the year over its full range.
Within Costa Rica, collected from January to March, and June to July.

HABITAT PREFERENCE. — This species seems to prefer woodlands. It has been collected in dry
forest, tropical rainforest and riverine vegetation. It has also been collected with the aid of light and
Malaise traps.

Nusalala ghioi Monserrat, 2000
(Figs. 518-520)

DIAGNOSIS. — A medium to large-sized brown species. Fore- and mid-tibiae with two dark
preapical spots. Forewing (Fig. 518) wide, subtriangular and somewhat acute at apex. Forewing com-
pletely dark variegated, with hexagonal cells between gradates and medial and outer gradates darker
than inner (forewing length 10-12 mm, hind wing length 9-11 mm). Male ectoproct and internal gen-
italia (Figs. 519-520).

GEOGRAPHICAL DISTRIBUTION. — A widely distributed species in Central America (Mexico,
Guatemala, Honduras, and Costa Rica). Within Costa Rica it has been collected in ALAJUELA:
Zarcero, and SAN JOSE: San Gerardo and Iraza.

ALTITUDINAL DISTRIBUTION. — Probably orophilous, 1520 to 3420 m. In Costa Rica, from
1000 to 2500 m.

ADULT FLIGHT PERIOD. — Collected from February to August, and November to December.
Costa Rican records are for February, April, May, and December.

HABITAT PREFERENCE. — This species appears to prefer forests. Collected in wet woodlands
and mixed tropical forest. Also collected with lights and Malaise traps.

Nusalala irrebita (Navas, 1929)
(Figs. 521-523)

DIAGNOsIS. — A large, pale brown species. Fore- and mid-femora and tibiae with two dark
preapical spots. Forewing (Fig. 521) widely subtriangular with somewhat acute apex, forewings pale
brown to dark variegated, with hexagonal cells between gradates and easy to distinguish due to two se-
ries of cells in its costal field (forewing length 8—10 mm, hind wing length 6-8 mm). Male ectoproct
and internal genitalia (Figs. 522-523).

GEOGRAPHICAL DISTRIBUTION. — Central America, rarely and only recorded from Mexico,
Honduras, El Salvador, Nicaragua, Costa Rica, and Panama. Within Costa Rica it has been collected
in PUNTARENAS: Monteverde; and SAN JOSE: Escazu.

ALTITUDINAL DISTRIBUTION. — 1300 to 1600 m in Costa Rica.

ADULT FLIGHT PERIOD. — Collected during almost the entire year (January, February, April to
September). Within Costa Rica collected in January, May, and September.

HABITAT PREFERENCE. — Rarely collected. Usually on vegetation in mixed forest and wet
woodlands. Also collected at lights.

Nusalala marginata Navas, 1926
This species was described from Costa Rica and subsequently reported from Mexico. Unfortu-

nately, from the original description it is not possible to assign this species to any known taxon in the
region and this species must be currently considered a nomen dubium.

248 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES
Volume 53, No. 12

Nusalala tessellata (Gerstaecker, 1888)
(Figs. 524-526)

DIAGNOSIS. — A medium-sized, pale brown species. Fore- and mid-tibiae with two dark
preapical spots. Wings (Fig. 524) subtriangular and somewhat acute at apex, pale brown, forewing
with darker variegated pattern, with hexagonal cells among gradates and darker external gradates
(forewing length 7-9 mm, hind wing length 6-7 mm). Male ectoproct and internal genitalia (Figs.
525-526).

GEOGRAPHICAL DISTRIBUTION. — A widely distributed species, broadly recorded in Central
America (Mexico, Guatemala, Honduras, Panama, Dominica, Virgin Islands, and Puerto Rico) and
South America (Colombia, Venezuela, Ecuador, Peru, Bolivia, Brazil, Paraguay, and Argentina).
Previously unknown from Costa Rica, where it has been amply collected in ALAJUELA: Upala;
Irazi; Zarcero; GUANACASTE: Cajias; Rio Naranjo; Santa Cecilia; San Lorenzo; and SAN JOSE:
Escazu.

ALTITUDINAL DISTRIBUTION. — Sea level to 1900 m; in Costa Rica from 400 to 1700 m.

ADULT FLIGHT PERIOD. — This species has been collected throughout the year. Within Costa
Rica, collecting records appear bimodal from January to May, and August to December, being espe-
cially abundant from February to March, and November to December.

HABITAT PREFERENCE. — This appears to be a generalist species. Collected in open and mixed
forest and fields, wet woodlands, savannas, on citrus, coffee, grasses and riverine vegetation. It has
also been collected at dusk and with the aid of light and Malaise traps.

Nusalala uncata Kimmins, 1937
(Figs. 527-529)

DIAGNOSIS. — A medium to large-sized, brown species. Fore- and mid-femora with a basal dark
spot and fore- and mid-tibiae with two dark preapical spots. Wings (Fig. 527) wide, subtriangular and
somewhat acute at apex, forewing darker than other species, fully dark variegated, with three dark
bands along the gradates and a longitudinal dark stripe joining them. Hind wings paler with dark exter-
nal gradates (forewing length 7-9 mm, hind wing length 6—8 mm). Male ectoproct and internal geni-
talia (Figs. 528-529).

GEOGRAPHICAL DISTRIBUTION. — Known only from Costa Rica and Panama. Within Costa
Rica it has been collected in CARTAGO: Quebrada Segunda; GUANACASTE; PUNTARENAS:
Las Alturas; Monteverde; and SAN JOSE: San Isidro del General; Escazu; Irazu.

ALTITUDINAL DISTRIBUTION. — Usually found at mid-elevation (610-2135 m). In Costa Rica
mostly found from 1100 to 1520 m and in Panama from 610 to 910 m.

ADULT FLIGHT PERIOD. — February, May to July, and September to October.

HABITAT PREFERENCE. — This species appears associated with moist woodlands. It has been
collected in wet forests and mixed tropical forest. It is attracted to lights.

Nusalala unguicaudata Monserrat, 2000
(Figs. 530-532)

DIAGNOSIS. — A medium-sized, pale brown species. Forefemora with two preapical dark spots
and fore- and mid-tibiae with two dark preapical spots. Forewing (Fig. 530) very elongated and with
acute apex, pale brown, forewing somewhat darkly variegated, with elongated cells between gradates
and dark margining on medial and outer gradates (forewing length 8-10 mm, hind wing length
7-8 mm). Male ectoproct and internal genitalia (Figs. 531—532).

PENNY: LACEWINGS OF COSTA RICA 249

GEOGRAPHICAL DISTRIBUTION. — Known only from Central America (Mexico, Guatemala, and
Costa Rica), where it has been collected in GUANACASTE: Rio Naranjo; ALAJUELA: Upala; and
PUNTARENAS: Las Alturas.

ALTITUDINAL DISTRIBUTION. — Elevation ca.1500 m.

ADULT FLIGHT PERIOD. — March, June, August, October, and Deceinber. Within Costa Rica
collected only in the latter part of the year from August, October, and December.

HABITAT PREFERENCE. — Unknown. Mostly collected with light traps.

Sympherobius Banks, 1904

This genus contains almost 60 valid species, widely distributed in Europe, western, northern and
central Asia, throughout the Americas, and northern, eastern, and southern Africa. The species from
America, north of Mexico, were revised by Oswald (1988) who also presented a world list of species.

Some of the previously described species from Central and South America are poorly known, and
probably new synonymies will be discovered. Our knowledge of Central American species 1s still in-
complete and new distribution records will certainly appear. Some of the species recorded here were
previously unknown from Central America, especially Costa Rica, where four species have now been
recorded.

KEY TO COSTA RICAN SYMPHEROBIUS

Note: Other species which may subsequently be found in Costa Rica include: S. perparvus
(McLachlan, 1869), S. barberi (Banks, 1903) and S. marginatus (Kimmins, 1928).

il3 Forewing without a crossvein connecting Ry,<; with R, or R,,, (Fig.534) ........---.--. S. arizonicus
I’. Forewing with a crossvein connecting Ry; with R, or R,,3 (Figs. 533, 536

2(1'). Forewing ovoid, with a dark spot basally in the subcostal space, outer gradate series especially darker than others,

and with a crossvein between R,,, and R,,, (Fig-536) .....----- 22+ eee eee ee eee S. subcostalis
2’. Forewing elongate, without a dark spot on the basal subcostal space; outer gradate series not especially darker than
other crossveins, and with a crossvein between Ry, 5 and R, (Pigs49339935)'2 ohare era ee See 3

3(2'). Longitudinal veins in forewing nearly concolorous brown, without alternating dark-hyaline segments (Fig. 535)
es eet TEN eM on chs) ey we. Se eye EY sled SRY Seon Be eS Gee S. similis
3’. _ Longitudinal veins in forewing with alternating dark-hyaline segments .................. S. angustus

Sympherobius angustus (Banks, 1904)
fie. 533)

DIAGNOSIS. — A small and variable pale brown species with brown antennae and forewings pale
and diffusely variegated, with darker stripe on Cu;; longitudinal veins alternating dark and hyaline,
and bearing a crossvein between fork of Ry.; and R, (Fig. 533). Forewing length 4.5—6.5 mm, hind
wing length 3.3—-5.3 mm. Body and genital morphology in Oswald (1988).

GEOGRAPHICAL DISTRIBUTION. — This species is known from Canada (British Columbia),
western U.S.A. (AZ, CA, NM, CO, ID, NV, OR, SD, TX, UT) and Mexico (Coahuila, Distrito Fed-
eral, Durango, Guanajuato, Jalisco, Michoacan, Nuevo Leon, Oaxaca). In Central America, it has
only been collected in Costa Rica: ALAJUELA: Zarcero.

ALTITUDINAL DISTRIBUTION. — 1100-2900 m. Within Costa Rica, it is known from 1700 m.

ADULT FLIGHT PERIOD. — From April to August, and October. Within Costa Rica it has been
collected in April and May.

250 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES
Volume 53, No. 12

HABITAT PREFERENCE. — This species appears to be associated with montane forests. They are
mainly collected at light traps.

Sympherobius arizonicus Banks, 1911
(Figs. 534)

DIAGNOSIS. — This is a small, pale brown species with dark brown antennae and brown varie-
gated forewings, with darker crossveins and without a crossvein between R4.; and R>.3 or Ry (Fig.
534). Forewing length 2.54.5 mm, hind wing length 3.2—5.2 mm. Body and genital morphology in
Oswald (1988).

GEOGRAPHICAL DISTRIBUTION. — Known from U.S.A. (AZ, CA) and Mexico (Baja California,
Oaxaca, Puebla, and Sonora). In Central America this species is only known from Costa Rica, where it
has been collected at: GUANACASTE: Canas Experimental Station (14 km SW Canas); and Santa
Rosa National Park.

ALTITUDINAL DISTRIBUTION. — Generally found at low elevations (30—1630 m). Within Costa
Rica it has been collected at 10 m.

ADULT FLIGHT PERIOD. — All months of the year. Within Costa Rica collection dates are Janu-
ary and March through September.

HABITAT PREFERENCE. — Unknown. Usually collected at light traps.

Sympherobius similis Carpenter, 1940
(Fi1g4535)

DIAGNOSIS. — A small, pale brown species with dark brown antennal flagellum and paler scape
and pedicel. Forewings alternating pale and dark variegations, with darker crossveins and bearing a
crossvein between R,,< and R,. Longitudinal veins almost uniformly dark brown and bearing a dark
oblique stripe on Cu, (Fig. 535). Forewing length 3.34.3 mm, hind wing length 2.7-3.7 mm. Body
and genital morphology in Oswald (1988).

GEOGRAPHICAL DISTRIBUTION. — Previously known only from U.S.A. (AZ), it is herein re-
corded for the first time from Mexico (Chiapas, Michoacan, Morelos, Nuevo Leon, and Veracruz) and
Central America (Costa Rica and Panama). Within Costa Rica this species has been collected from
GUANACASTE: Rio Naranjo; and HEREDIA: Santo Domingo.

ALTITUDINAL DISTRIBUTION. — In Panama at 1300 m. The Rio Naranjo locality is near sea
level, while Santo Domingo de Heredia is in the central valley near San José at above 1000 m.

ADULT FLIGHT PERIOD. — January, April, June, and July. Within Costa Rica it has been col-
lected in January and April.

HABITAT PREFERENCE. — Unknown. Mostly collected at light traps.

REMARKS. — Previous records of Sympherobius miranda (Navas, 1920) in Costa Rica
(Monserrat 1990) pertain to this species.

Sympherobius subcostalis Monserrat, 1990
(Fig. 536)

DIAGNOsIS. — A small pale brown species with pale flagellum, but darker brown on segments 10
to 15. Forewings oval, the external crossveins darker, with a dark spot on the basal subcostal
crossvein, and a crossvein between R4,5 and R>,3 (Fig. 536). Forewing length 3.5—-4.5 mm, hind wing
length 3.0-4.0 mm. Body and genital morphology in Monserrat (1990).

PENNY: LACEWINGS OF COSTA RICA Za

GEOGRAPHICAL DISTRIBUTION. — Known from Mexico (Chiapas, Jalisco, Veracruz, and
Yucatan) and Central America (Nicaragua and Costa Rica). It has also been recorded from Colombia.
Within Costa Rica it has been collected in abundance in GUANACASTE: Canas; and Rio Naranjo.

ALTITUDINAL DISTRIBUTION. — 230 to 630 m. The Canas and Rio Naranjo localities are at low
elevations in flat to rolling hills of agricultural lands.

ADULT FLIGHT PERIOD. — Outside Costa Rica from January, May, June, and August. Within
Costa Rica from January to June, September, October, and December.

HABITAT PREFERENCE. — Unknown. Mostly collected at light traps and in forests. Some speci-
mens have been collected on citrus and Clerodendrum.

FAMILY MANTISPIDAE

by
Kevin Hoffman

Adults of the family Mantispidae are sometimes called mantisflies because of the combination of
an elongate prothorax and raptorial prolegs, which causes adults to resemble small praying mantises.
The family is divided into four subfamilies, three of which occur in Costa Rica. Adult mantispids are
mostly predators of small insects, however, some species also feed on plant exudates and at floral and
extrafloral nectaries. Many species are attracted to lights at night, while others are primarily diurnal
and can be found on or near flowers. Larvae are predators of sedentary arthropods and spider eggs. Co-
coons are constructed in association with the larval food source by using silk spun through an anal
spinneret. The exarate pupae exit cocoons by chewing holes in them with decticious mandibles and
may wander some distance before adult eclosion. Eighteen species are known from Costa Rica. An
additional species is included in the following treatment because it is distributed in the region and
therefore will most likely be collected in Costa Rica in the future.

KEY TO GENERA OF NORTH AND CENTRAL AMERICAN MANTISPIDAE

I. Prolegs each with two pretarsal claws and a median arolium (Figs. 538-540); head with vertex domed in frontal
wlan (Fige, SAR=SS Oh Sethe barred: gases amranet ee Nem Aiea Rien citi eesti Ss teeee ae Neale wet team Aiton Yc 2
is Prolegs each with one tarsal claw and no arolium (Fig. 541); head with vertex concave in frontal view (Fig. 551)
a :o foun ear ee Suplannhy. Mantispinne sy tse eae Geer nes tee ake PLA. WE Ere a eee Oe Cee

2(1). Pronotum at least twice as long as high in lateral view (Fig. 555); basal tarsomere of protarsus without dentiform
process (Fig. 540); wing with pterostigmal spot semicircular, entirely dark (Fig. 544)... 2... ....-.-.--

eeeeSubianmiyCalomantispinaes - 94 56 5 2 a) se ee ee ee ae ey ane Nolima
2. Pronotum only slightly longer than high in lateral view (Figs. 553, 554); basal tarsomere of protarsus with

lanceolate process (Figs. 538, 539); wing with pterostigmal spot elongate, dark with middle pale (Figs. 542, 543). .

Pee Subiannlysymohrasinae tabwewacn. es 22 eG coe = oe ie oA EN ee OR ee Ral eek 3
BGs) eee toOlemurwithisno-pasal spine (big. 558). 2 4-4. 265 2-s 526 ae 22 eee ee ee se Plega
Bf. Brotemumwithoutsub-pasalispine(Pigs539)). 44.425 55.4 55 5556.0 4. 9952s ae a = Trichoscelia

4(1'). Antennal flagellomeres at mid-length of flagellum each three or more times as wide as long in anterior view
(Figs. 563-567); mesoscutal furrow obsolete (Figs. 558-562); forewing with anterior half pale amber to dark

[Divo (riveree oy eg Goo SUETGeS to GMs Suse er er Gme cao Omeeeoe cect ona) Geer oma) poecr uec race Climaciella
4’. Antennal flagellomeres at mid-length of flagellum less than three times as wide as long in anterior view; mesoscutal
furrow conspicuous (Figs. 555-557); forewing with anterior half hyaline ........-..-.-.-.-+--. S
5(4'). Pronotum with prominent setae over entire length in lateral view (Figs. 608,609) ........-.-.-.-.-- 6

ae Pronotum with scattered fine setae in lateral view, most setae at anterior and posterior ends (Figs. 606,607)... 9

252 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES
Volume 53, No. 12

6(5). Pronotum with most setae arising from distinct bumps (Fig. 609); forewing with Sc distal to sc-ra at 30° angle to

RAG(PIG. S89 591) oo comisiel ping tins youl 21> Sey hte eit Hoa er eteou out boa ae eet ie a Zeugomantispa
6. Pronotum with most setae arising flush with pronotal surface (Fig. 608); forewing with Sc distal to sc-ra at 40—50°
angle to RA (EIS SGD). oo ns So Os ee ae tiga eS 6 ee es rr q/.

7(6'). | Pronotum 4.4—S.0 times as long as wide at maculae; forewing with membrane around |ra-rp and 2ra-rp amber to
light brown; male abdomen with long setae emanating from pores on tergites !V-V anterolaterally; pseudopenis
broadly triangular and as wide as pseudopenal membrane; female spermatheca with proximal section distinctly

wider than medial'section. sc 2 a tee) 3) Gps pieces Ges ee ee Ste cay er 3 Xeromantispa
Ps Pronotum 5.5—9.0 times as long as wide at maculae; forewing with membrane around Ira-rp and 2ra-rp hyaline;
male abdomen without long setae emanating from pores on tergites [V-V; pseudopenis spinelike in posterior view;
female spermatheca with proximal section as wide as, or narrower than, medial section ............. 8
8(7'). Forewing with amber infuscation posterior to 1MP (Fig. 569); body mostly darkred ...... Haematomantispa
8. Forewing without amber infuscation posterior to IMP (Figs. 586-588); body mostly a mixture of yellow and
DIOWD) ene tee cee On ee aes oe pe ey i Gy Ree atu beac) acs hae Leptomantispa

9(5'). Head with inverted V-shaped mark directly beneath antennae (Fig. 552); wing apex infuscated (Fig. 568); body

coloration mottledcamoutlage-like<u.-. sc) huts shear elicite ci & idee be efeale coe Buyda
oF Head with either longitudinal or transverse stripe beneath antennae (Fig. 579); wing apex unmarked; body color-
ATOM VATIOG) eee cr ue oe wie 2) os 6 Re ey des ae cake. oe tune hs St ee a 10

10(9'). Head mostly brown above antennae (Fig. 582); mesonotum with paired longitudinal yellow stripes laterally (Fig.
583), mostly brown anterior to mesoscutal furrow; forewing with basal third of cell IRA hyaline (Fig. 572)
drodtubober ie felt bad tey bch uel GaPade. |) cote cee ers Gert ode bee Dicromantispa
10’. Head yellow with transverse brown stripes above and beneath antennae (Fig. 551); mesonotum with paired yellow
stripes angled from wing bases to scutellum (Fig. 556), almost entirely yellow anterior to mesoscutal furrow;
forewing usually with basal third of cell IRA amber (Fig.570) .................... Entanoneura

SUBFAMILY CALOMANTISPINAE

This subfamily contains two genera, one of which occurs in Costa Rica. Little is known of their
life histories. The only rearings of a calomantispine are of No/ima pinal Rehn from the southwestern
United States (MacLeod and Redborg 1982). Larvae fed on Coleoptera larvae and pupae, Lepidoptera
pupae, aculeate Hymenoptera larvae, Diptera pupae, and the eggs and paralyzed adults of spiders. The
immature stages were not described, but all three larval instars were fully ambulatory and conceivably
could feed on several different prey items during the larval life span.

Nolima Navas, 1914
Nolima contains seven species, one of which occurs 1n Costa Rica.

Nolima infensus Navas, 1924
(Figs. 540, 544, 550, 555)

DIAGNOsIS. — Nolima infensus can be separated from other Costa Rican mantispids by the dark,
semicircular, pterostigmal spot on each wing. In addition, this species has each protarsus with two
pretarsal claws and a median arolium and lacks a dentiform process on the basal tarsomere.

GEOGRAPHICAL DISTRIBUTION. — This species is known only from Costa Rica. It was described
by Navas (1924) from an unspecified locality, and has since been collected in the province of San José.
The specific locality is as follows: SAN JOSE: Escazu.

ALTITUDINAL DISTRIBUTION. — Unknown.

ADULT FLIGHT PERIOD. — July.

HABITAT PREFERENCE. — Unknown.

PENNY: LACEWINGS OF COSTA RICA 253.

SUBFAMILY MANTISPINAE

Fifteen species are presently known to occur in Costa Rica. The following information on the
subfamily is excerpted from the work by Hoffman (1992). Adults have been reared exclusively from
the egg sacs of spiders. Larvae feed on eggs by piercing the chorion and draining the contents, and
have been noted to feed on neonate spiderlings as well. Larvae have three instars and are
hypermetamorphic with the first instar campodeiform and the last two scarabaeiform. Female
mantispines lay clutches of approximately 200 to 2000 individually stalked eggs on objects such as
leaves, twigs, branches, wooden structures, and iron poles. Following eclosion, first-instar
mantispines must locate spider egg sacs and gain access to the eggs within. The two general methods
used by larvae to gain access to eggs are either to locate and enter preexisting egg sacs or to locate and
board female spiders and enter egg sacs as they are deposited. Different mantispine species may use
one of these two strategies exclusively or either strategy facultatively. Larvae aboard spiders gener-
ally are found either wrapped around the pedicel, inside the book lung openings, or attached to the
membranous area between the edge of the carapace and the base of the legs; such larvae have survived
for months apparently by feeding on spider haemolymph and can be considered true ectoparasites at
this stage in their life cycle.

Buyda Navas, 1926

This genus contains two species occurring from Honduras to northern Argentina, one of which is
known from Costa Rica. The immature stages are unknown.

Buyda phthisica (Gerstaecker, 1885) [new combination]
(Figs. 552, 557, 568, 606)

DIAGNOSIS. — Buyda phthisica can be separated from other Costa Rican mantispines by the
combination of an inverted V-shaped mark directly beneath the antennae, the infuscated wing tips,
and the mottled, camouflage-like body coloration.

GEOGRAPHICAL DISTRIBUTION. — This species occurs from Honduras to northern Argentina.
Within Costa Rica, specific localities are as follows: CARTAGO: Turrialba; near Turrialba;
HEREDIA: Chilamante; La Selva Biological Station, 3 km S Puerto Viejo, 10°26'N, 84°01'W; Finca
La Selva, 4 km SE Puerto Viejo de Sarapiqui; LIMON: Hacienda Tapezco, 29 air km W Tortuguero,
10°30'N, 83°47'W;: Rio Banano, 16 km WSW Bomba, 9.888°N, 83.167° W; Rio Uatsi, 8 km (air) W
Bribri, 9.62°N, 82.90°W; PUNTARENAS: Osa Peninsula, 2.9 km (1.8 mi) W Rincon.

ALTITUDINAL DISTRIBUTION. — 40-150 m.

ADULT FLIGHT PERIOD. — February to June, and October.

HABITAT PREFERENCE. — Unknown.

Climaciella Enderlein, 1910

This genus contains nine species found from southern Canada to northern Argentina, including
the Caribbean islands of Cuba, Hispaniola, and Puerto Rico. Three species occur in Costa Rica. All of
the species are somewhat wasplike in appearance, and can be easily distinguished from other Costa
Rican mantispines by wings that have the anterior half amber to dark brown. Adults of Climaciella
brunnea (Say), which occurs in Costa Rica, are diurnal and commonly found on various flowers. The
immature stages of C. brunnea have been associated with four species of wolf spiders (Lycosidae),
and first instars have been described.

PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES
Volume 53, No. 12

i)
Nn
Ss

KEY TO COSTA RICAN CLIMACIELLA

1. Pronotum distinctly bent ventrad at mid-length in lateral view (Fig. 605); profemur with basal third reddish with
black spot medially; wings with anterior three-fourths dark brown and posterior fourth hyaline to very light amber

(Hig SAG) roe, clea cece tases ive ety eke eee, Se atacel monsimshae est Teh fo cty net reps dharas poo rn C. obtusa
I’. Pronotum straight in lateral view (Figs. 603, 604); profemur without basal third reddish or with black spot medially;

wings with anterior half brown to amber and posterior fourth light amber to amber (Figs. 545,547) ........ 2
2(1'). Pronotum with distinct hump dorsomedially in lateral view (Fig. 604). ................... C. porosa
2'. Pronotum without distinct hump dorsomedially in lateral view (Fig. 603), at most with slightly raised, flattened

KERION Re ee eon es Ti i cl ds ween Gee ys) 8. ANS cue elven aan We ee C. brunnea

Climaciella brunnea (Say, 1824)
(Figs. 545, 558-560, 563-565, 603)

DIAGNOsIS. — Climaciella brunnea can be separated from other Costa Rican mantispines by the
combination of the pronotum being straight in lateral view and the wings with the anterior half brown
to amber and posterior fourth light amber to amber. Climaciella brunnea is a polychromatic species
whose different color forms have been presumed to be mimics of various species of polistine wasps
(Hymenoptera: Vespidae: Polistinae). Opler (1981) described the five color forms occurring in Costa
Rica and listed the possible wasp models.

GEOGRAPHICAL DISTRIBUTION. — This species occurs from southern Canada south to Costa
Rica. Within Costa Rica specific localities are as follows: CARTAGO: Turrialba; GUANACASTE:
Comelco Ranch, 8 km NW Bagaces; Hacienda La Pacifica, 4 km NW Canas; Hacienda La Pacifica,
5 km N Canas; La Pacifica, nr. Canas; Taboga, 9.7 km S, 9.7 km W Canas, 10°19’N, 85°09’W; 14 km
S Cafias; Headquarters of Parque Nacional Santa Rosa, NW of Liberia; SAN JOSE: 10 km N San
José.

ALTITUDINAL DISTRIBUTION. — 300—900 m.

ADULT FLIGHT PERIOD. — Every month except March.

HABITAT PREFERENCE. — Unknown.

Climaciella obtusa Hoffman, new species
(Figs. 546, 561, 566, 605)

(non) Euclimacia semihyalina (Lepeletier and Audinet-Serville), Navas 1927 (in part): 39 (Costa Rica: Cartago).
(non) Climaciella semihyalina (Lepeletier and Audinet-Serville), Hughes-Schrader 1969:110, 122 (female
karyotype, Panama: Canal Zone), fig. 45 (female chromosomes), table 3 (chromosome numbers).

DIAGNOSIS. — Separated from other Costa Rican mantispines by the combination of the
pronotum being distinctly bent ventrad at mid-length in lateral view and the wings with the anterior
three-fourths dark brown.

MALE COLORATION. — Head black (Fig. 566), posterior margin of eyes and vertex yellow;
mouthparts dark reddish brown, mandible reddish brown distally, lacinia and galea yellow-brown to
reddish brown, palps dark reddish brown with apex yellow-brown; antenna with scape brown, pedicel
and flagellum dark reddish brown; eyes metallic silver to light purple. Prothorax with pronotum and
membranes black, pronotum very dark reddish brown ventrally; pleural sclerites and membranes very
dark reddish brown, appearing blackish, sternum reddish brown; leg with coxa very dark reddish
brown, distal fifth orange-red, sulcus sometimes yellow, trochanter reddish brown, femur orange-red
proximad of major spine with black spots medially on each side, major spine dark reddish brown, re-
mainder black, tibia very dark reddish brown, tarsus with tarsomere I dark reddish brown, tarsomeres
II-V orange. Pterothorax black, metathoracic posterodorsal lobes margined with yellow posteriorly;

PENNY: LACEWINGS OF COSTA RICA 255

legs each with coxa very dark reddish brown, nearly black, trochanter and femur very dark reddish
brown, tibia and tarsus dark reddish brown; forewing with membrane mostly dark amber (Fig. 546),
posterior 0.2—0.3 hyaline, pterostigma and asperous region of 1AP brown, veins dark brown; hind
wing similar to forewing. Abdomen black, pleura I-II yellow medially, tergite and sternite II usually
with posterior margins narrowly yellow.

FEMALE COLORATION. — Same as for males.

THORAX. — Pronotum 2.2—2.5 times as long as wide at maculae in dorsal view (Fig. 561):
pterothoracic scutella with combined 4-8 pores on each side, pretarsal claws with 5—6 teeth each;
males with both wings collectively on each side with combined 14—17 veins leaving RP posteriorly,
females with 17—18 veins; forewing lengths of males 15.8—24.0 mm, of females 17.0—-24.2 mm.

MALE PREGENITAL ABDOMINAL APPARATUS. — Tergites IV-V each with paired patches of
18-36 small circular pores total in two transverse rows on each side anterolaterally, rows converging
near midline on each side, pores with small conical collars internally, tergites V-VI with
intersegmental membrane expanded into very slightly bilobed pocket extending anteriorly one-third
length of tergite V, tergite V 1.5 times as long as VI.

MALE TERMINALIA. — Ectoprocts each ovoid in dorsal view, apex rounded, ventromedial lobe
flattened dorsoventrally, lightly sclerotized ventrally, abruptly truncate along inner margin, with
50-65 short thickened setae dorsally and posteriorly; sternite [X pentagonal in ventral view, scooplike
in lateral view, deepest near anterior margin, posterior margin produced into short,
posteroventrally-directed, rounded lobe medially; gonarcus with medial third obliquely flattened
anteroposteriorly, flattened region convex in lateral view and transversely ovoid in posterior view,
median lobe spinelike, curved anteriorly, 2—3 times as long as wide at base; gonocoxites each as wide
distally as basally in lateral view, distal fourth slightly bent dorsolaterally, apex narrowly rounded;
mediuncus with anterior fourth slightly higher than remainder in lateral view, ventral surface laterally
expanded into circular to oval plate extending 0.2—0.5 length of mediuncus, apex forked in posterior
view; pseudopenal membrane broadly triangular, lightly sclerotized; hypomeres present as paired,
small, granulate, circular to elliptical sclerites; pseudopenis sclerotized, spinelike in posterior view,
slightly longer than pseudopenal membrane, flattened anteroposteriorly, apex broadly rounded to
truncate, one-half or more as wide as base.

FEMALE TERMINALIA. — Ectoprocts as long to slightly longer than gonocoxites; sternite VIII 1.5
times as long medially as laterally, most of medial 0.2—0.3 separated posteriorly from sternite by nar-
row membranous region, medial section transversely elliptical with posterior margin slightly con-
cave, posterior margin of remainder of sternite truncate; bursa lightly sclerotized; spermatheca with
proximal, medial, and distal sections equal in width, diverticulae present as low transverse ridge,
proximal section with first bend nestled within first spiral of medial section, medial section spiralling
anteriorly for 1.5 loops before abruptly bending posteriorly, distal section spiralling posteriorly for
1.0—-1.5 loops and intertwining with medial section, fertilization canal duct curved, fertilization canal
truncate basad of apex.

GEOGRAPHICAL DISTRIBUTION. — Costa Rica (CARTAGO); Panama (Canal Zone); Colombia
(Magdalena); and Ecuador (Los Rios, Pichincha).

ALTITUDINAL DISTRIBUTION. — Elevation 900 m within Costa Rica.

ADULT FLIGHT PERIOD. — No information available for Costa Rica. Elsewhere during March,
May, June, August, and October.

ETYMOLOGY. — Name is derived from the Latin adjective obtusa, meaning blunt or dull, and is
in reference to the abruptly blunted apex of the male pseudopenis.

TYPE MATERIAL. — Climaciella obtusa holotype male (Florida State Collection of Arthropods,
Gainesville [FSCA]), six paratype males, eight paratype females, and two paratypes of undetermined
gender (American Museum of Natural History, New York [AMNH], Carnegie Museum of Natural
History, Pittsburgh, PA [CMNH], FSCA, Deutsches Entomologisches Institut, Eberswalde-Finow,

256 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES
Volume 53, No. 12

Germany [DEIC], Museum of Comparative Zoology, Harvard University, Cambridge, MA [MCZC],
Museu de Zoologia, Universidade de Sao Paulo, Brazil [MZSP}, San Diego Natural History Museum
[SDMC], University of Minnesota Insect Collection [UMIM], US National Museum of Natural His-
tory, Washington [USNM]). Holotype label data: “RIO PALENQUE, ECUADOR/ LOS RIOS
PROV. — C. DODSON / 197,” “503,” “HOLOTYPE MALE / Climaciella / obtusa Hoffman / det:
K. M. Hoffman 1992.” Condition of holotype: color excellent, wings unspread, forewing length
20.0 mm. Paratypes have the following label data associated with them: COSTA RICA: Cartago
Prov., Cartago, N. Banks, | undet. (MCZC); Cartago Prov., Turrialba, Heyne, Berlin-Wilm., 1 male
(DEIC); same data except 900 m, IV, 1 male (DEIC): Cartago Prov., Turrialba, 4., E. A. Bottcher,
Berlin C. 2, | female (DEIC); PANAMA: Canal Zone, Barro Colorado Island, 1—9-V-1964, W. D.
and S. S. Duckworth, | female (USNM); same locality, XI-1965, 399, Dr. Britsky, 1 female (MZSP);
same locality, 6-X-1956, Ch. Gregoire, a 1538, 1 female (USNM); same locality, 19-HI-1936,
Gertsch, Lutz and Wood, | male (AMNH); Canal Zone, 9 km NW Gamboa, 10—12-VIII-1975, E. M.
and J. L. Fisher, 1 female (SDMC); COLOMBIA: Magdalena Div., Bonda, (77 m), June, H. H.
Smith, Carn. Mus. Acc. 1999, 1 undet. (MCZC); Magdalena Div., Don Diego (31 m), May, H. H.
Smith, Carn. Mus. Acc. 1999, 1 female (CMNH); ECUADOR: Los Rios Prov., Rio Palenque, 197, C.
Dodson 503, | male, | female (FSCA); same locality, 12-VI-1974, J. Longino 503, 1 male (FSCA);
same locality, 12-VIII-1975, J. Longino, | female (FSCA); Pichincha Prov., Hotel Tinalandia, 12 km
E Santo Domingo de los Colorados, 23-V-1985, Clausen and Cervenka, | male (UMIM).

Climaciella porosa Hoffman, new species
(Figs. 547, 562, 567, 604)

DIAGNOSIS. — Climaciella porosa can be separated from other Costa Rican mantispines by the
combination of the pronotum bearing a distinct hump dorsomedially in lateral view and by the wings
having the anterior half brown to amber.

MALE COLORATION. — Head with vertex and frons dark brown (Fig. 567), remainder yel-
low-brown, labrum with dark brown spot proximomedially; mandibles dark brown with yel-
low-brown proximomedially, other mouthparts yellow-brown, palps brown with distal segment
yellow; antenna with scape yellow-brown with brown posterodistally, pedicel dark brown, flagellum
with basal half light brown anteriorly, remainder light red with several apical flagellomeres pale yel-
low; eyes metallic purple. Prothorax with pronotum dark brown anterior of maculae, remainder
brown dorsally, yellow-brown laterally and ventrally, posterior half with paired thin dark brown sinu-
ous stripes laterally, posterodorsal membrane dark brown, posteroventral membrane pale yellow;
pleural sclerites and membrane dark brown, sternum brown; leg with coxa brown, trochanter yel-
low-brown, femur dark reddish brown anterolaterally, reddish brown posterolaterally, yellow-brown
proximad of major spine ventrally, tibia dark brown anterolaterally and distally, remainder and tarsus
yellow to yellow-brown. Pterothorax with nota dark brown, pleura reddish brown, anterior membrane
yellow-brown; legs each with coxa reddish brown, trochanter reddish brown, femur light brown to
brown, tibia yellow-brown to light brown with apex brown, tarsus yellow-brown; forewing with ante-
rior half amber (Fig. 547), posterior margin light amber, remainder hyaline, pterostigma and asperous
region of 1AP brown, veins brown; hind wing similar to forewing. Abdomen dark brown, pleura I-II
pale yellow.

FEMALE COLORATION. — Same as for male, except for the following: head with vertex, frons,
clypeus, and labrum very dark brown, remainder brown; mandibles very dark reddish brown, other
mouthparts dark brown; antenna with scape and pedicel dark brown, flagellum very dark brown with
several apical flagellomeres brown; pronotum darker than that of male; proleg with coxa dark brown,
femur dark reddish brown, spines reddish brown distally, tibia dark brown, tarsus with tarsomere I

PENNY: LACEWINGS OF COSTA RICA 2S

brown; pterothoracic legs each with trochanter yellow-brown proximally, brown distally, femur and
tibia brown; abdomen yellow-brown to brown.

THORAX. — Pronotum 3.3—3.7 times as long as wide at maculae in dorsal view (Fig. 562), dis-
tinct hump dorsomedially (Fig. 604); pterothoracic scutella with combined 3-4 pores on each side,
pretarsal claws with five teeth each; male with both wings collectively on each side with combined
14-15 veins leaving RP posteriorly, female with 16 veins; forewing length of male 12.7 mm, of fe-
male 14.7 mm.

MALE PREGENITAL ABDOMINAL APPARATUS. — Tergites IV-V each with paired patches of
137—160 small circular pores total in 10-12 transverse rows occupying entire tergite on each side lat-
erally, rows continuous across midline, pores each surrounded by dark microtrichia externally and
with small conical collars internally, tergites V- VI with intersegmental membrane not noticeably ex-
panded into pocket, tergite V 1.25 times as long as VI.

MALE TERMINALIA. — Ectoprocts each ovoid to crescent-shaped in dorsal view, lobelike poste-
rior to ventromedial lobe and as long as wide, apex rounded, ventromedial lobe flattened
dorsoventrally, lightly sclerotized ventrally, abruptly truncate along inner margin, with 60—70 short
thickened setae dorsally and posteriorly; tergite [X with ventral third with posterior margin concave;
sternite [X triangular in ventral view, shallowly scooplike in lateral view, deepest near middle, poste-
rior margin produced into short, posteroventrally-directed, rounded lobe medially; gonarcus with me-
dial third obliquely flattened anteroposteriorly, flattened region convex in lateral view and
transversely ovoid in posterior view, median lobe spinelike, curved anteriorly, twice as long as wide at
base; gonocoxites each as wide distally as basally in lateral view, distal fourth bent dorsolaterally,
apex narrowly rounded, separate flattened extension at three-fourths length directed medially up onto
gonarcal membrane for short distance, as long as distal fourth of gonocoxite, apex pointed; mediuncus
as high anteriorly as posteriorly in lateral view, ventral surface laterally expanded into small circular
plate extending 0.2—0.4 length of mediuncus, apex truncate in posterior view; pseudopenal membrane
broadly triangular, lightly sclerotized; hypomeres present as paired linear longitudinal series of 8-10
very small granules; pseudopenis sclerotized, spinelike in posterior view, distinctly shorter than
pseudopenal membrane, flattened anteroposteriorly.

FEMALE TERMINALIA. — Ectoprocts as long as gonocoxites; sternite VIII 1.5 times as long me-
dially as laterally, most of medial fifth separated posteriorly from sternite by narrow membranous re-
gion, medial section transversely elliptical with posterior margin slightly concave, posterior margin
of remainder of sternite truncate; bursa lightly sclerotized; spermatheca with proximal, medial, and
distal sections equal in width, diverticulae absent, area at diverticulae slightly concave with small
flanges laterally, proximal section with first bend nestled within first spiral of medial section, medial
section spiralling anteriorly for one loop before abruptly bending posteriorly, distal section bending
anteriorly and then posteriorly, and curving along the right side of medial section, fertilization canal
duct slightly curved basally, slightly twisted medially, fertilization canal sloping basad of apex.

GEOGRAPHICAL DISTRIBUTION. — Costa Rica (HEREDIA).

ALTITUDINAL DISTRIBUTION. — 50 m.

ADULT FLIGHT PERIOD. — May and June.

ETYMOLOGY. — Name is derived from the Latin adjective porosa, meaning full of pores, and is
in reference to the pores of males, which occupy the full length of abdominal tergites IV and V in
males.

TYPE MATERIAL. — Holotype male (Henry A. Hespenheide Collection [CHAH]) and paratype
female (Clemson University Arthropod Collection [CUAC]). Holotype label data: “CosTA RICA:
Heredia / Pr: La Selva Biol. Sta. / 3 km S Pt. Viejo / 10°26'N, 84°01'W,” “19.v.1990 / H. A.
Hespenheide,” “HOLOTYPE male / Climaciella porosa / Hoffman / det: K. M. Hoffman 1992.” Con-
dition of holotype: color good, right antennal flagellum absent; right wings spread, forewing length
12.7 mm; abdomen detached, cleared, in glycerin-filled microvial attached to pin beneath labels. La-

tN
Nn
CO

PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES
Volume 53, No. 12

bel data associated with paratype: COSTA RICA: Heredia Prov., Est. Biol. La Selva, Quebrada Sura,
10.437°N, 84.010°W, el. 50 m, 20—21-VI-1986, Holzenthal, Heyn, Armitage, | female (CUAC).

Dicromantispa Hoffman, new genus

TYPE SPECIES. — Mantispa sayi Banks (1897) by present designation.

DESCRIPTION. — Head with shortest distance between eyes narrower than width of one eye at
ventral margin of antennal sockets in frontal view, antennal flagellum at mid-length with
flagellomeres 1—2 times as wide as long in anterior view. Pronotum straight in lateral view (Fig. 607),
3.37.3 times as long as wide at maculae in dorsal view, dorsal surface mostly glabrous, with some
scattered fine setae, setal bases not elevated. Pterothorax with mesoscutal furrow prominent,
pterothoracic scutella with 0-7 pores total on each side; legs each with tarsomere I nearly as long as or
longer than tarsomeres II-V combined, pretarsal claws with 3—7 teeth each; both wings collectively on
each side with combined 8-32 veins leaving RP posteriorly, forewing length 7.0—28.9 mm,
pterostigma with sc-ra at 0.5—0.8 length of 3RA (Fig. 571), Sc distal to sc-ra at 40—-60° angle to RA,
3RA with one c-ra crossvein distal to SC, AA and AP; simple, 1 AP with posterior 0.3—0.5 asperous;
hind wing with CuA strongly angled toward AA, cu-aa crossvein either very short or absent with CuA
briefly fused with AA. Male abdomen with tergites IV-VI each with paired patches of 2—17 circular
pores total in two transverse rows on each side anterolaterally, pores recessed in shallow circular to el-
liptical pits, tergite V 1.3 times as long as VI. Male terminalia with each ectoproct roughly ovoid in
dorsal view and rounded distal to ventromedial lobe, ventromedial lobe strongly sclerotized and
deeply cleft to form dorsal plate and finger-like ventral lobe, dorsal plate with 240 short thickened
setae on dorsal margin, ventral lobe with 3—15 short, thickened setae at apex anteriorly; sternite 1X
semicircular to trapezoidal in ventral view; gonarcus with paired, anteriorly-projected apodemes
sometimes present dorsolaterally, median lobe absent or small, anteroposteriorly flattened, and scale-
like; gonarcal membrane unmodified or with small spinules medially; each gonocoxite lightly
sclerotized, variously widened distally, apex rounded; mediuncus with anterior third 1.5—3.0 times as
high as remainder in lateral view, anterior third sometimes excavated ventrally, apex forked;
pseudopenal base membranous; hypomeres granular to elongate; pseudopenis sclerotized, spinelike;
hypandrium internum triangular in ventral view, shallow and nearly straight in lateral view. Female
terminalia with ectoprocts as long as, or longer than, gonocoxites; sternite VIII entire, transverse
sclerotized ridge usually present immediately dorsal of and partially hidden by sternite VIII; bursa
with entrance membranous to lightly sclerotized, remainder membranous to lightly sclerotized;
spermatheca with proximal and medial sections nearly equal in width, distal section wider than medial
section, diverticulae present as transverse ridge with paired small bumps or as medially-directed lobe,
medial portion with 0.5—3.5 coils, fertilization canal duct bent at 0.3—0.5 length, fertilization canal
slightly curved to C-shaped, sloped or truncate basad of apex.

CHROMOSOME COMPLEMENT. — Diploid chromosomal complement of 22 consisting of 20
autosomes and 2 sex chromosomes, based on D. sayi. Sex chromosomes of males are XY and those of
females are XX.

IMMATURE STAGES. — Immature stages of D. sayi have been described (Hoffman and
Brushwein 1992). The stalked eggs are oval with a caplike micropyle and are typical for mantispines.
First instars are diagnosed as follows: head capsule brown, thoracic terga with paired brown rectangu-
lar to triangular markings, abdominal terga I-VIII white anteriorly, brown posteriorly, [X-X brown;
mandibles untoothed, antennal scape cylindrical, longer than wide; legs with tarsus wider than long,
unguitractor process up to half width of tarsus and as long as tarsal claw; abdominal segments I-VI
with setae D4, SD3, and L2 at most barely reaching posterior margins of segments. Second and third
instars are diagnosed as follows: legs with tarsus subquadrate, claw present; second instar with ab-
dominal segment X shorter than [X, third instar with spigot less than twice as long as base. Pupae are

PENNY: LACEWINGS OF COSTA RICA 259

diagnosed as follows: abdominal tergite IV with setae very short and not reaching above dorsal hooks,
VIII with setae in two transverse rows laterally.

BIoLoGy. — Adults have been captured at lights and on vegetation. Immatures of D. sayi have
been associated with 46 species of primarily hunting spiders in 14 families (Hoffman and Brushwein
1992), and D. gracilis has been reared from egg sacs of Trechalea manausensis Carico (Araneae:
Pisauridae) (Carico et al. 1985). The vast majority of recorded spider hosts for members of this genus
are wandering hunting spiders. First instars of D. sayi board spiders and are known to be facultative
boarder/penetrators.

GEOGRAPHICAL DISTRIBUTION. — Southern Canada south to Argentina, including the Bahama
Islands, Cuba, and Puerto Rico.

ETYMOLOGY. — A combination of the Greek adjective dicro, meaning forked, and mantispa as
coined by Illiger (1798), which is a combination of the Greek noun mantis, meaning soothsayer or
prophet, and the suffix pa, which is derived either from the Greek para, meaning near or beside, or the
Latin par, meaning equal. The name ts in reference to the deeply cleft, ventromedial lobes on the male
ectoprocts. Gender is feminine.

DISCUSSION. — This genus contains eight species, four of which have been collected in Costa
Rica. Members of this genus can be distinguished by the combination of the head being mostly brown
above antennae, the presence of only a few scattered setae on the pronotum in lateral view with most
setae at the anterior and posterior ends, paired longitudinal yellow stripes on the mesonotum laterally,
and the mesonotum anterior to the mesoscutal furrow being brown. Of the Costa Rican species,
immatures of D. sayi (Banks) have been associated with 46 species of spiders in 14 families, and D.
gracilis (Erichson) has been reared from egg sacs of Trechalea amazonica F. O. Pickard-Cambridge
(Trechaleidae). All larval instars and pupae of D. sayi have been described.

KEY TO COSTA RICAN DICROMANTISPA

1. Forewing with Sc yellow to yellow-brown throughout its length, distinctly lighter than costal crossveins; cell 1M

Eg Or SR eo ict Sr ale OBS a yal SPY papeme rte Givers tpotaltlye: fered ye D. gracilis
1’. Forewing with Sc brown basally, at least on basal half, essentially same color as costal crossveins; cell 1M mostly

ainbemomal least rduiaimancleand Sco space amber (BIS, SVL), 62. 4 ee ss ee ee ee 2,
2(1'). Antennae tricolored, with basal 2/3 brown, next 1/6 yellow, and apical 1/6 dark brown ............ D. sayi
SP ELC IIS AG Nene lve ans OrOW le semede te Mane iewts in ees Matra epee Rt (SP SIG. baeicedon a Saeus See cer SE mac taek Vee 3

3(2'). Head above antennae with yellow marks forming a broad, procurved band (Fig. 583); pterothorax with anterior

MESO remy el OweCdORSally se seet ui we eye See eo ay ew oe es ce ce oe ss i oe ys as eee D. synapsis
3’. Head above antennae with yellow marks narrow, not forming a broad band across head (Fig. 580); pterothorax with
Aci OMe mbOrinc now GOIsallyens sc <5 es 2 te See A eee ee ce eee fe Sis nee emo se D. debilis

Dicromantispa debilis (Gerstaecker, 1888) [new combination]
(Figs. 574, 580)

DIAGNOSIS. — Separated from other Costa Rican mantispines by the combination of a pronotum
with only a few scattered setae in lateral view with most setae at the anterior and posterior ends, paired
longitudinal yellow stripes on the mesonotum laterally, the forewing with Sc brown basally and es-
sentially the same color as the costal crossveins, antennae entirely dark brown, the head above the an-
tennae with thin, curved, yellow marks forming medially interrupted circle (Fig. 580), and the
pterothorax with the anterior membrane brown dorsally.

GEOGRAPHICAL DISTRIBUTION. — Costa Rica south to Bolivia. Within Costa Rica it is only
known from: HEREDIA: La Selva Biological Station, 3 km S Puerto Viejo, 10°26'N, 84°01'W.

ALTITUDINAL DISTRIBUTION. — Unknown.

260 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES
Volume 53, No. 12

ADULT FLIGHT PERIOD. — June and July in Costa Rica.
HABITAT PREFERENCE. — Unknown.

Dicromantispa gracilis (Erichson, 1839) [mew combination]
@igss5724575;.581))

GEOGRAPHICAL DISTRIBUTION. — Costa Rica south to Argentina. Within Costa Rica specific lo-
calities are as follows: GUANACASTE: 3 mi NW Canas (La Pacifica), PUNTARENAS: Golfito;
4.5 km E Golfito; Rio Palmare Ferry Crossing, nr. Brujo, Pan American Hwy.

ALTITUDINAL DISTRIBUTION. — 150 m.

ADULT FLIGHT PERIOD. — July and September.

HABITAT PREFERENCE. — Unknown.

Dicromantispa sayi (Banks, 1897) [new combination|
(Figs. 571, 576, 582, 607)

DIAGNOSIS. — Separated from other Costa Rican mantispines by the combination of the
pronotum with only a few scattered setae in lateral view with most setae at the anterior and posterior
ends, paired longitudinal yellow stripes on the mesonotum laterally, the forewing with Sc brown ba-
sally and essentially the same color as the costal crossveins, head above the antennae with paired
(sometimes fused), small, comma-shaped, yellow marks medially (Fig. 582), and the tricolored an-
tennae, with the basal 2/3 brown, next 1/6 yellow, and apical 1/6 dark brown.

GEOGRAPHICAL DISTRIBUTION. — Most of the United States south to Panama, including the
Bahama Islands, Cuba, and Puerto Rico. Within Costa Rica specific localities are as follows:
GUANACASTE: Hacienda Tempisquito (Pelon de la Altura), 1 km NE of km 265, rt 1, 10.84°N,
85.56°W; Hacienda La Pacifica; La Pacifica, nr. Canas; La Pacifica, 4 km NW Canas; 14 km S Canas;
Parque Nacional Santa Rosa; PUNTARENAS: Golfito.

ALTITUDINAL DISTRIBUTION. — 100-300 m.

ADULT FLIGHT PERIOD. — Every month except August and September.

HABITAT PREFERENCE. — Unknown.

Dicromantispa synapsis Hoffman, new species
(Higs¥5 7325774583)

DIAGNOSIS. — Separated from other Costa Rican mantispines by the combination of the
pronotum with only a few scattered setae in lateral view with most setae at the anterior and posterior
ends, paired longitudinal yellow stripes on the mesonotum laterally, the forewing with Sc brown ba-
sally and essentially the same color as the costal crossveins, antennae entirely dark brown, the head
above the antennae with yellow marks forming a broad, procurved band, and the pterostigma with the
anterior membrane yellow dorsally.

MALE COLORATION. — Head yellow (Fig. 577), vertex mostly covered by inverted brown trian-
gular mark, brown extending between antennal bases, slightly procurved transverse yellow stripe
above antennae, ends of stripe obscuring lateral margins of brown region, sometimes a yellow stripe
along the midline connecting with transverse stripe to form inverted Y-shaped mark, lateral margins
sometimes indented at mid-length, a thin transverse brown stripe sometimes extending to ocular mar-

PENNY: LACEWINGS OF COSTA RICA 261

gins beneath antennal bases, anterior half of frons sometimes with brown triangular mark medially,
brown stripe along midline from between antennae, from midfrons, or from clypeus and onto labrum;
mandibles orange-brown, sometimes pale yellow distomedially, other mouthparts yellow with palps
yellow-brown; antenna with scape yellow anteriorly, brown posteriorly, pedicel brown, flagellum
dark brown; eyes metallic silver. Prothorax with pronotum yellow-brown to light brown, usually yel-
low ventrally, paired wide diffuse brown stripes from anterolateral corners to maculae (Fig. 583),
maculae brown, paired wide diffuse brown stripes from mid-length to posterolateral margins,
posterodorsal membrane entirely yellow to yellow-brown or yellow anteriorly, brown posteriorly,
posteroventral membranes yellow; cervical sclerite brown dorsally, yellow ventrally, episternum,
sternum, and membrane yellow; leg with coxa yellow with light brown distad of sulcus posteriorly,
trochanter yellow-brown, femur yellow-brown anterolaterally with dark reddish brown medially
from apex to base of major spine, yellow to yellow-brown posterolaterally with base of spine row dark
orange-brown to brown, dorsal surface yellow-brown, spines orange-brown, tibia yellow with apex
reddish brown anteriorly, tarsus with tarsomere I reddish brown, remainder yellow to yellow-brown.
Pterothorax with nota brown to dark reddish brown, mesonotum with narrow longitudinal yellow
stripes at one-third width on each side, stripes mostly obscure on metanotum, pleura yellow,
mesopreepisternum, mesokatepisternum, and anterior margin of mesanepisternum brown some-
times, mesopreepisternum yellow with paired small diffuse reddish brown spots laterally sometimes;
legs each with coxa yellow, mesocoxa sometimes brown, remainder of leg segments yellow to yel-
low-brown; forewing membrane hyaline (Fig. 573), Sc space basad of radial triangle, radial triangle,
1M, most of 1Cuand 1 AA light brown, remainder of Sc space very light amber to opaque, pterostigma
dark reddish brown to dark brown, sometimes with basal fourth yellow, 1AP with asperous region
pale yellow, most veins brown, fused C+Sc sometimes Sc inside pterostigma, RA basad of sc-ra dor-
sally, proximal half of CuA dorsally, AA yellow except at apex, AP, AP>, and wing margin basad of
AA yellow; hind wing similar to forewing, Sc space entirely very light amber to opaque, | A hyaline,
pterostigma entirely dark reddish brown to dark brown, Sc basad of pterostigma yellow except at base,
RA brown except along 2RA. Abdomen with tergites yellow with broad dark reddish brown stripe
along midline, stripe widening posteriorly on each segment, ectoprocts dark brown; pleura I-II black,
remainder yellow to dark reddish brown dorsally, dark brown ventrally and widening posteriorly on
each segment; sternites yellow-brown or yellow with brown posteromedially, discolored.

FEMALE COLORATION. — Same as for males, except for the following: antennae sometimes with
scape entirely brown; procoxa brown distad of sulcus posterolaterally and posteriorly; prothorax with
pronotum entirely brown anterior of maculae dorsally; sometimes with cervical sclerite entirely yel-
low; procoxa sometimes light brown distad of sulcus laterally, profemur entirely dark reddish brown
anterolaterally, posterior midline yellow proximad of spine row; pteropleura with margins of anterior
membrane dark brown, mesopreepisternum, katepisterna, katepimera, and most of mesanepisternum
dark brown, metanepisternum and anepimera sometimes brown posteriorly, coxae brown to dark
brown or mesocoxa brown anteriorly and metacoxa brown anterolaterally, trochanter usually light
brown medially; abdominal tergites very dark brown instead of dark reddish brown, pleura entirely
dark brown to black, sternites yellow I-VI with posterior margins dark reddish brown, I-IV sometimes
entirely yellow with VI dark brown, VII entirely dark brown, VIII light brown to brown, ectoprocts
and gonocoxites dark reddish brown to yellow.

MALE PREGENITAL ABDOMINAL APPARATUS. — Tergires II-V with medial sixth of posterior
margins sclerotized and slightly concave, IV-VI each with paired patches of 3—8 circular pores total in
two transverse rows on each side anterolaterally, pores with very small conical collars internally,
pores recessed in shallow circular to elliptical pits, tergite V 1.3 times longer than VI.

MALE TERMINALIA. — Ectoprocts each ovoid in dorsal view, apex rounded, ventromedial lobe
entirely sclerotized, laterally-flattened, platelike, with thin dorsally-curved lobe from posteroventral
corner, roughly triangular in lateral view, posterior margin longer than others, margins relatively

262 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES
Volume 53, No. 12

straight, anteroventral corner broadly rounded, dorsal apex rounded with 2-5 short thickened setae,
posteroventral lobe rounded laterally with 4—12 shorter thickened setae at apex anteriorly; sternite IX
somewhat quadrate in ventral view, posterior margin slightly produced medially and rounded api-
cally; gonarcus without median lobe, gonarcus produced anteriorly with paired short
dorsoventrally-flattened lobes; gonarcal membrane with subquadrate patch of golden yellow, pros-
trate, anteriorly-directed spinules medially, posterior corners extended posteriorly into thin lobes;
gonocoxites each sclerotized, distal half twice as wide as remainder, apex rounded, extended dorsally
onto gonarcal membrane and joining with posterolateral lobes of spinule patch; mediuncus with ante-
rior half ovoid in lateral view, 2—3 times as high as remainder, apex forked; pseudopenal membrane
triangular in posterior view, lightly sclerotized; hypomeres teardrop-shaped, widest ventrally and ir-
regularly granulate; pseudopenis sclerotized, spinelike in posterior view, longer than height of
pseudopenal membrane, flattened laterally, bladelike in lateral view.

FEMALE TERMINALIA. — Ectoprocts longer than gonocoxites; sternite VIII as long medially as
laterally, posterior margin broadly rounded to truncate medially, very slightly concave at midline,
dorsal surface lightly sclerotized laterad of midline, membrane immediately posterior to dorsal sur-
face lightly sclerotized, less so at midline, and collapsing against dorsal surface of sternite VIII to
form pocket distinctly shorter than length of sternite; membrane between sternite VIII and bursa with
transverse, rectangular, sclerotized patch of pointed to rounded microspines medially, patch four
times as wide as long, three-fourth width of sternite VIII, distinctly convex; bursa lightly sclerotized
to membranous; spermatheca with proximal and medial sections equal in width, distal section twice as
wide as proximal, proximal section with first bend laterad of medial section, sinuous between first and
second bends, diverticula present as low ridge or absent, medial section with two bends, fertilization
canal duct darkened at base, sinuous, fertilization canal truncate basad of apex.

THORAX. — Pronotum 5.6—7.3 times as long as wide at maculae in dorsal view; pterothoracic
scutella with combined 3-7 pores on each side, pretarsal claws with 5—6 teeth each, males with both
wings collectively on each side with combined 13—15 veins leaving RP posteriorly, females with
11—16 veins, forewing lengths of males 9.3—13.7 mm, of females 9.8—15.8 mm.

GEOGRAPHICAL DISTRIBUTION. — Costa Rica (CARTAGO, LIMON); Panama (Bocas del
Toro, Panama); French Guiana (Cayenne); Brazil (Rio de Janeiro, Santa Catarina).

ADULT FLIGHT PERIOD. — In Costa Rica March and May. They have been collected during Feb-
ruary, May, June, and from October to December in other countries.

ALTITUDINAL DISTRIBUTION. — Elevation 60 m in Costa Rica.

ETYMOLOGY. — Name is derived from the Greek noun synapsis, meaning a connection, union,
or junction, and is in reference to the unique articulation of the male gonocoxites dorsally with the
patch of spinules on the gonarcal membrane.

TYPE MATERIAL. — Holotype male (Peabody Museum of Natural History, Yale University,
New Haven, CT [PMNH)]), three paratype males, ten paratype females, and one paratype of undeter-
mined gender (AMNH, CMNH, CUAC, FSCA, LSUC, MCZC, PMNH, SDMC). Holotype label
data: “Brasilien / Nova Teutonia / 27° 11'B, 52°23'L/ Fritz Plaumann / XI-1962 / 300-500 m” [date
and altitude are written vertically, with date along right margin and altitude along left margin].
“HOLOTYPE male / Dicromantispa synapsis / Hoffman / Det. K. M. Hoffman 1991.” Condition of
holotype: color excellent, forewing length 13.8 mm, abdomen detached, cleared, in glycerin-filled
microvial attached to pin beneath labels. Paratypes have the following label data. COSTA RICA:
Cartago Prov., Turrialba-C.A.T.I.E., 28-3 1-V-1987, E. Giesbert, 1 female (FSCA); Limon Prov., Rio
Uatsi, 8 km (air) W. Bribri, 9.62°N, 89.90°W, el. 60 m, 25-III-1987, Holzenthal, Hamilton, Heyn, 3
females (CUAC); PANAMA: Bocas del Toro Prov., 10 km NE Fortuna Dam, 1—2-VI-1986, E.
Giesbert, | female (FSCA); Panama Prov., 10 km N El Llano, 16—22-V-1987, E. Giesbert, 1 male
(FSCA); FRENCH GUIANA: Cayenne Dept., Hwy N2 to Regina, 67 km S of Cayenne, 4-VI-1986,
E.G. Riley and D. A. Rider, 1 female (LSUC); BRAZIL: PARA: Oriximina, Rio Trombetas,

PENNY: LACEWINGS OF COSTA RICA 263

ALCOA Miner., km 22, 28-XI-1982, N. Penny, 1 female (SDMC); SANTA CATARINA: Corupa
(Hansa Humboldt), H-1946, A. Maller, Coll., Frank Johnson, Donor, 1 female (AMNH); Nova
Teutonia, 27°11’B, 52°23’L, 300-500 m, X-1962, Fritz Plaumann, 1 female (PMNH); Nova
Teutonia, 27° 11'B, 52°23'L, 30-XI-1939, Fritz Plaumann, | male (MCZC); RIO DE JANEIRO: Rio
de Janeiro, Dec., Acc. No. 2966, H. H. Smith, | female (CMNH); Rio de Janeiro, Dec., H. Smith, 1
undet. (MCZC); Rio, Nov., H. Smith, | male (MCZC).

Entanoneura Enderlein, 1910

This genus contains six species occurring from southern Mexico south to northern Argentina, one
of which has been collected in Costa Rica. The immature stages are unknown.

Entanoneura batesella (Westwood, 1867) [new combination]
fies. 5515556, 570, 602)

DIAGNOSIS. — Separated from other Costa Rican mantispines by the combination of transverse
brown stripes above and beneath the antennae, the forewing usually with the basal third of cell IRA
amber, the mesonotum yellow anterior to the mesoscutal furrow and with paired yellow stripes angled
from wing bases to the scutellum on the mesonotum.

GEOGRAPHICAL DISTRIBUTION. — Southern Mexico south to Brazil. Within Costa Rica specific
localities are as follows: CARTAGO: Turrialba; 15 km NE of Turrialba; HEREDIA: La Selva Bio-
logical Station, 3 km S Puerto Viejo, 10°26'N, 84°01'W; La Selva Biological Station, nr. Puerto
Viejo de Sarapiqui; 1 km S Pt. Vieja; LIMON: plain of Limon near Las Mercedes, Farm Hamburg at
Reventazon.

ALTITUDINAL DISTRIBUTION. — Unknown.

ADULT FLIGHT PERIOD. — February, April, June, August, and December.

HABITAT PREFERENCE. — Unknown.

REMARKS. — Entanoneura limbata (Gerstaecker, 1885), which has been reported from Costa
Rica, is a new synonym of Entanoneura batesella.

Haematomantispa Hoffman, new genus

TYPE SPECIES. — Mantispa nubeculosa Navas, 1933 by present designation.

264 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES
Volume 53, No. 12

anteroposteriorly-flattened lobe medially or with toothlike median lobe, gonarcus with anterior mar-
gin produced dorsally behind lobe into anteroposteriorly-flattened flange either medially or at
dorsolateral corners; gonarcal membrane without spinules; gonocoxites each wider distally than ba-
sally in lateral view, distal half convex, abruptly narrowed to point apically; mediuncus notched
anteroventrally in lateral view, nearly uniform height throughout, apex forked in posterior view;
pseudopenal membrane lightly granulate; hypomeres present as paired small, darkened, circular
sclerites or absent; pseudopenis sclerotized, setalike, apex curved anteriorly; hypandrium internum
triangular in ventral view, shallow and with posterior point slightly curved dorsad in lateral view. Fe-
male terminalia with ectoprocts shorter or longer than gonocoxites; sternite VIII entire, longer medi-
ally than laterally; bursa with entrance lightly sclerotized, remainder lightly sclerotized to
membranous; spermatheca with proximal section as wide as most of medial section, medial section
gradually widened after first bend, distal section 2—3 times as wide as proximal section, proximal sec-
tion sinuous between bends, diverticulae present as one very small bump or as paired small bumps,
medial section with two bends or 1.5 coils, fertilization canal duct sinuous, fertilization canal truncate
basad of apex.

CHROMOSOME COMPLEMENT. — Unknown.

IMMATURE STAGES. — Unknown.

BIOLOGY. — Adults have been captured at lights.

GEOGRAPHICAL DISTRIBUTION. — Costa Rica and Panama.

ETYMOLOGY. — A combination of the Greek adjective haemato, meaning blood red, and
mantispa as coined by Illiger (1798), which is a combination of the Greek noun mantis, meaning
soothsayer or prophet, and the suffix pa, which is derived either from the Greek para, meaning near or
beside, or the Latin par, meaning equal. The name is in reference to the striking red coloration of the
included species. Gender is feminine.

Haematomantispa nubeculosa (Navas, 1933) [new combination|
(Figs. 569, 578, 584)

DIAGNOSIS. — Separated most easily from other Costa Rican mantispines by the combination of
an amber infuscation posterior to cell 1 MP in the forewing and the mostly dark red body coloration.

GEOGRAPHICAL DISTRIBUTION. — This species occurs in Costa Rica and Panama. Within Costa
Rica specific localities are as follows: ALAJUELA: Reserva Forestal San Ramon, Rio Lorencito and
tribs., 10.216°N, 84.607°W; CARTAGO: Turrialba; near Turrialba; HEREDIA: La Selva Biological
Station, 3 km S Puerto Viejo, 10°26'N, 81°01'W; Puerto Viejo de Sarapiqui; LIMON: Reserva Biol.
Hitoy-Cerere, Rio Cerere, 9.671°N, 83.028°W; Rio Banano, 16 km WSW Bomba, 9.888°N,
83.167° W; Rio Uatsi, 8 km (air) W Bribri, 9.62°N, 82.90°W; near Rita (Hamburg, at Reventazon);
locality not found: Paso Marcos; Pta. Uvita.

ALTITUDINAL DISTRIBUTION. — 40-1000 m.

ADULT FLIGHT PERIOD. — January to August, and November.

HABITAT PREFERENCE. — Unknown.

Leptomantispa Hoffman, new genus

TYPE SPECIES. — Mantispilla pulchella Banks, 1912 by present designation.

PENNY: LACEWINGS OF COSTA RICA 265

with combined 2-6 pores on each side; legs each with tarsomere I longer than tarsomeres II-V com-
bined, pretarsal claws with 3—5 teeth each; both wings collectively on each side with combined 6—14
veins leaving RP posteriorly, forewing length 5.2-14.5 mm, pterostigma with sc-ra at less than
mid-length of 3RA, Sc distal to sc-ra at 40—50° angle to RA (Fig. 586), 3RA with one c-ra crossvein
distal to Sc, AA and AP; simple, 1AP asperous along AP»; hind wing with CuA strongly angled to-
ward AA and fused with AA for varied distance, 2cu-m sometimes absent, CuA sometimes with
3cu-m absent and with terminal branch fused with MP, AA often fused with wing margin prior to fu-
sion with CuA. Male abdomen with tergites III-IV each with or without pores along posterior margin,
1-17 pores on each side of tergite when present, tergites [V-V each with pores anterolaterally, 6—40
pores on each side of tergite, tergite V 1.7—3.0 times as long as VI, intersegmental membrane between
tergites V-VI usually invaginated into single-lobed pocket extending anteriorly 0.2—0.4 length of ter-
gite V. Male terminalia with each ectoprct ovoid to crescent-shaped in dorsal view, apex rounded,
sclerotized ventrad of ventromedial lobe, ventromedial lobe sometimes somewhat flattened
dorsoventrally, abruptly truncate along inner margin, with 8—35 short thickened setae; tergite 1X
sometimes with ventral fourth concave anteriorly; sternite [X pentagonal to triangular in ventral view,
posterior margin rounded medially; gonarcus with median lobe spinelike to lobelike, slightly curved
anteriorly, |—4 times as long as wide at mid-length; each gonocoxite up to three times as wide distally
as basally in lateral view, apex rounded or pointed; mediuncus with anterior 0.2—0.4 up to three times
as high as remainder in lateral view, apex forked in posterior view; pseudopenal membrane broadly
ovoid to rectangular in posterior view, with dorsally-projecting microspines or granules laterally and
often medially; hypomeres present as paired small circular sclerites or granules; pseudopenis
sclerotized; spinelike in posterior view, less than twice as long as pseudopenal membrane;
hypandrium internum triangular in ventral view, shallow and nearly straight in lateral view. Female
terminalia with ectoprocts shorter or longer than gonocoxites; sternite VIII 1—2 times as long medially
as laterally; bursa with entrance lightly sclerotized, remainder membranous to lightly sclerotized;
spermatheca with proximal section as wide as, or wider than, medial section, distal section wider than
proximal, diverticula present as thin transverse rod or as medially-directed lobe from inner margin,
proximal section with first bend right laterad of medial section, medial section twisting anteriorly for
2-4 bends, distal section C-shaped in ventral view, fertilization canal duct curved to sinuous, fertiliza-
tion canal sloping basad of apex.

CHROMOSOME COMPLEMENT. — Diploid chromosome complement of 22 consisting of 20
autosomes and 2 sex chromosomes, based on L. pulchella (Hughes-Schrader, 1969). Sex chromo-
somes of males are XY and those of females are XX.

IMMATURE STAGES. — Larvae and pupae of L. pulchella have been described (Hoffman and
Brushwein 1992). First instars are diagnosed as follows: head capsule pale white, tergites with
bracketlike brown markings laterally; head with setae S1 and S4 surpassing lateral margin of head,
mandibles with three teeth on inner margin medially, antennal scape cylindrical, slightly longer than
wide; legs with tarsus wider than long, unguitractor process nearly as wide as tarsus and nearly twice
as long as tarsal claw, empodium five times as wide distally as basally; abdominal segments I-VI with
setae D4, SD3, and L2 distinctly surpassing posterior margins of segments. Second and third instars
are diagnosed by the following: legs with tarsus peglike, claw absent; abdominal segment X shorter
than IX, second instar with segment X broadly conical apically, third instar with spigot less than twice
as long as base. Pupae are diagnosed by the following: abdominal tergites III-IV each with setae 2—3
times as long as height of dorsal hooks in lateral view, tergite VIII with setae in one transverse row lat-
erally.

BIOLOGY. — Adults have been collected at lights and on foliage. Immatures of L. pulchella have
been associated with 21 spider species in 8 families (Hoffman and Brushwein 1992), and are spider
boarders.

266 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES
Volume 53, No. 12

ETYMOLOGY. — A combination of the Greek adjective /epto, meaning frail or scaled, and
mantispa as coined by Illiger (1798), which is a combination of the Greek noun mantis, meaning
soothsayer or prophet, and the suffix pa, which is derived either from the Greek para, meaning near or
beside, or the Latin par, meaning equal. The name refers both to the relatively small size and frail body
construction and to the dorsally-projecting scalelike microspines on the male pseudopenal mem-
brane. Gender is feminine.

REMARKS. — This genus contains eight species occurring from southwestern Canada south to
northern Argentina, including islands of the Greater and Lesser Antilles. Two species are known from
Costa Rica, and a third has a distribution which brackets the country and is therefore included in the
summary below. Members of this genus can be distinguished by the combination of prominent setae
over the entire length of the pronotum in lateral view with most setae arising flush with the pronotal
surface, the forewing lacking an amber infuscation posterior to cell 1 MP, and with Sc distal to sc-ra at
a 40—50° angle to RA in the forewing. The immatures of Leptomantispa pulchella (Banks), which oc-
curs in Costa Rica, have been associated with 21 species of spiders in 8 families, and all three larval in-
stars and the pupa have been described.

KEY TO COSTA RICAN LEPTOMANTISPA

1. Mesoscutellum brown medially, yellow laterally (Fig. 585); mesofemur with longitudinal brown stripe ventrally;

forewing with subcostal space hyaline, pterostigma lightred......................2.. L. pulchella
1’. Mesoscutellum entirely yellow (Figs. 597, 598); mesofemur without longitudinal brown stripe ventrally; forewing

with subcostal space distinctly amber, pterostigma reddish brown to dark brown... ................ 2
2(1’). Pronotum entirely dark reddish brown to dark brown (Fig. 597); forewing with pterostigma dark brown . . L. chaos
2’. Pronotum yellow-brown to brown dorsally (Fig. 598); forewing with pterostigma reddish brown to brown L. nymphe

Leptomantispa chaos Hoffman, new species
(Figs: 587, 592, 597)

DIAGNOsIS. — Separated from other Costa Rican mantispines by the combination of prominent
setae over the entire length of the pronotum in lateral view with most setae arising flush with the
pronotal surface, pronotum entirely dark reddish brown to dark brown, mesoscutellum entirely yel-
low, forewing with subcostal space distinctly amber and the pterostigma dark brown.

MALE COLORATION. — Head yellow, vertex brown except along ocular margins, paired short
transverse yellow stripes somewhat posteriad of antennae and usually not meeting at midline, narrow
elliptical yellow spot above and between antennae, brown stripe along midline from slightly anteriad
of antennae and onto labrum, stripe often forked at antennae with ends curling around antennal bases
posteriorly, widened beneath antennae to form transverse brown stripe, clypeus usually with distal
margin brown; mandibles yellow basally, reddish brown distally, other mouthparts yellow, stipes,
prementum, and ligula yellow-brown to brown, palps light brown; antenna with scape yellow anteri-
orly, reddish brown posteriorly, pedicel reddish brown, flagellum dark brown; eyes metallic purple.
Prothorax with pronotum dark reddish brown to brown, posterodorsal membrane dark brown,
posteroventral membrane yellow with dark brown laterally; pleural sclerites and sternum brown,
membrane pale yellow; leg with coxa dull yellow, apex brown, trochanter yellow-brown, femur pale
yellow at base, ventral midline yellow basad of major spine, dark reddish brown anterolaterally, red-
dish brown posterolaterally, base of spine row pale yellow, tibia reddish brown, yellow-brown
posterolaterally, tarsus reddish brown. Pterothorax with nota brown, scutella yellow, yellow spot at
each wing base; pleura yellow, mottled with dark brown especially anteriorly on pleural sclerites, an-
terior membrane pale yellow with dark brown Y-shaped mark medially, sometimes variously spotted
with brown; legs each with coxa yellow, mottled with brown, trochanter brown, femur yellow-brown,

PENNY: LACEWINGS OF COSTA RICA 267

base brown, tibia and tarsus yellow-brown; forewing membrane hyaline, Sc space amber, pterostigma
dark brown, 1 AP with asperous region yellow-brown, most veins dark brown, AP, yellow basally;
hind wing similar to forewing, AA yellow except at apex. Abdomen with tergites dark brown, tergites
I-V with posterolateral corners yellow; pleura dark brown, spiracles and dorsal and ventral margins
yellow; sternites dark reddish brown to dark brown, paired oval yellow spots sublaterally, sternites
VI-IX with spots extending length of sternites.

FEMALE COLORATION. — Same as for males, except for the following: pterothorax with pleura
nearly entirely brown; abdomen with tergites entirely dark reddish brown, pleura with only ventral
margin yellow, sternites dark reddish brown with paired small yellow spots sublaterally on II-VI.

THORAX. — Pronotum 6.4—7.4 times as long as wide at maculae in dorsal view; pterothoracic
scutella with combined 24 pores on each side, pretarsal claws with four teeth each, males and fe-
males with both wings collectively on each side with combined 6-8 veins leaving RP posteriorly,
forewing lengths of males and females 6.8—9.5 mm.

MALE PREGENITAL ABDOMINAL APPARATUS. — Tergites HI-I'V each with paired patches of
3—17 small circular closely-packed pores total in one transverse row on posterior margin on each side,
IV-V each with paired patches of 14-40 small circular pores total closely packed into inverted
U-shaped band on each side anterolaterally, posterior arm of U-shaped band shorter than anterior arm
on tergite V and nearly absent on IV, tergites V-VI with intersegmental membrane invaginated into
single-lobed pocket extending anteriorly two-fifths length of tergite V, tergite V three times as long as
VI.

MALE TERMINALIA. — Ectoprocts each ovoid to crescent-shaped in dorsal view, apex rounded,
sclerotized ventrad of ventromedial lobe, ventromedial lobe sclerotized, flattened posteriorly,
abruptly truncate along inner margin, with 12—22 short thickened setae dorsally and posteriorly;
sternite |X pentagonal in ventral view, posterior margin broadly rounded medially; gonarcus with me-
dian lobe projecting from posterior surface, flattened posteriorly and projecting slightly posteriad
from gonarcus before curving dorsally, slightly curved anteriorly, lobelike in posterior view, 2—3
times as long as wide, apex rounded; gonocoxites each twice as wide distally as basally in lateral view,
apex narrowed and rounded; mediuncus with anterior third ovoid to semicircular in lateral view, 2—3
times as high as remainder, apex forked in posterior view; pseudopenal membrane ovoid in posterior
view, covered with golden yellow dorsally-projecting microspines; hypomeres present as paired
small granules; pseudopenis sclerotized, spinelike in posterior view, slightly curved posteriorly, as
long as pseudopenal membrane and longer than gonarcal median lobe.

FEMALE TERMINALIA. — Ectoprocts as long as gonocoxites; sternite VIII strongly curved poste-
riorly, slightly longer medially than laterally, medial fifth with posterior margin truncate to shallowly
concave; bursa with entrance lightly sclerotized, remainder membranous to lightly sclerotized;
spermatheca with proximal section as wide as medial section, distal section three times as wide as
proximal, diverticulae present as thin transverse medially-directed lobe from inner margin of
spermatheca, proximal section with first bend right laterad of medial section, medial section with 2—3
bends, fertilization canal duct sinuous to twisted, fertilization canal sloping basad of apex.

GEOGRAPHICAL DISTRIBUTION. — Guatemala (Alta Vera Paz); Panama (Canal Zone); Vene-
zuela (Aragua); French Guiana (Saint Laurent du Maroni); Brazil (Para). It has not been collected to
date in Costa Rica, but its distribution brackets the country and it will most likely be collected in the fu-
ture.

ALTITUDINAL DISTRIBUTION. — Unknown for Costa Rica.

ADULT FLIGHT PERIOD. — Unknown for Costa Rica. Adults have been captured in other coun-
tries during January, March, April, June, and July.

ETYMOLOGY. — Name derived from the Greek noun chaos, meaning utter confusion or disorder,
and is in reference to the rather difficult time I had deciding whether the specimens assigned to this
species were distinct from L. pulchella.

268 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES
Volume 53, No. 12

TyPE MATERIAL. — Holotype male (MCZC), 8 paratype males, and 4 paratype females
(CMNH, FMNH, John R. MacDonald Collection [JRMC], LSUC, MCZC). Holotype label data:
“PANAMA: Barro / Colorado Is. / Canal Zone,” “Robert G. Beard / 2 July 1967 / at U.V. light,”
“HOLOTYPE male / Leptomantispa chaos / Hoffman / det. K. M. Hoffman 1991.” Condition of
holotype: color good, wings spread, forewing length 9.5 mm, abdomen detached, cleared, in glyc-
erin-filled microvial attached to pin beneath labels. Paratypes have the following data associated with
them: GUATEMALA: Alta Vera Paz Dept., Larquin, elev. 1000 ft., 11-VI-1948, R. D. Mitchell,
Guatemala Zool. Exped. (1948), 1 female (FMNH); PANAMA: Canal Zone, Barro Colorado Island,
2-VII-1967, at U.V. light, Robert G. Beard, 2 females (MCZC); same data, 10-VII-196?, 1 female
(MCZC); same locality, 4-IV-1959, at light, #720, H. S. Dybas, 1 male (FMNH); Canal Zone, Ft. San
Lorenzo, 2-I-1983, E. Giesbert, | male (SDMC); Canal Zone, Gatun, Ridge Road, 29—30-III-1987,
J. R. MacDonald, 2 males (JRMC; VENEZUELA: Aragua St., Rancho Grande, 1100 m, 5-VII-1967,
R. W. Poole, 1 male (MCZC); same data, 20-VII-1967, 1 male (MCZC); FRENCH GUIANA: Saint
Laurent du Maron Dept., 13 km SSE of St. Laurent, 8-VI-1986, E. G. Riley and D. A. Rider, 1 male
(LSUC); BRAZIL: PARA: Tucurui, I-1979, M. Alvarenga, | male (CMNH).

Leptomantispa nymphe Hoffman, new species
(Figs. 588, 593, 598)

DIAGNOSIS. — Separated from other Costa Rican mantispines by the combination of prominent
setae over the entire length of the pronotum in lateral view with most setae arising flush with the
pronotal surface, pronotum yellow-brown to brown dorsally, mesoscutellum entirely yellow,
forewing with subcostal space distinctly amber and the pterostigma reddish brown to brown.

MALE COLORATION. — Head dull yellow, vertex brown posteriorly except along ocular mar-
gins, reddish brown spot above and between antennae medially, sometimes extended posteriorly
along midline, brown transverse stripe sometimes beneath antennae, stripe not surpassing antennal
bases laterally, frons with distal half dark brown medially, reddish brown stripe along midline from
frons and onto labrum; mandibles reddish brown, other mouthparts yellow to yellow-brown; antenna
with scape yellow, brown posterodistally, pedicel dark reddish brown, flagellum very dark reddish
brown; eyes metallic silver to purple. Prothorax with pronotum yellow-brown to brown dorsally,
anterolateral corners somewhat darker; posterodorsal membrane brown, posteroventral membrane
yellow; pleural sclerites and sternum yellow to light brown, membrane yellow; leg with coxa yellow,
trochanter light brown, femur light brown to light reddish brown, base of spine row yellow-brown,
tibia and tarsus brown. Pterothorax with nota brown, midline yellow, anterolateral corners of
mesonotum yellow, yellow spots at wing bases, scutella yellow; pleura yellow, anterior membrane
yellow medially, dark reddish brown marginally, all sclerites except metanepisternum and
metanepimeron partly brown; legs each with coxa and trochanter brown, femur, tibia, and tarsus dull
yellow; forewing with membrane hyaline, Sc space amber, pterostigma reddish brown to brown, 1 AP
with asperous region brown, most veins brown, RA pale yellow at base, most of AA and AP, pale yel-
low basally; hind wing similar to forewing, RA yellow for longer distance than in forewing, MP pale
yellow at base. Abdomen with tergites yellow, I-III mostly dark brown medially, HI-VIII reddish
brown posteriorly, ectoprocts brown; pleura yellow anteriorly, dark brown posteriorly, or almost en-
tirely dark brown; sternites yellow, I[V-VII reddish brown posteriorly.

FEMALE COLORATION. — Same as for males, except for the following: pronotum with
anterolateral surface of femur dark reddish brown medially; abdomen with tergites VI-IX mostly dark
brown, ectoprocts yellow, brown distally and along inner margin, pleura VI-VII entirely dark brown
to black, sternites VI-VII and gonocoxites dark brown.

THORAX. — Pronotum 6.7—8.1 times as long as wide at maculae in dorsal view; pterothoracic
scutella with combined 2-4 pores on each side, pretarsal claws with four teeth each; males with both

PENNY: LACEWINGS OF COSTA RICA 269

wings collectively on each side with combined 8—10 veins leaving RP posteriorly, females with 10
veins; forewing lengths of males 9.7—10.7 mm, of females 10.0—10.7 mm.

MALE PREGENITAL ABDOMINAL APPARATUS. — Tergites HI-I[V each with paired patches of
10-17 small circular pores total in one irregular transverse row on posterior margin on each side, rows
slightly angled anteriorly near midline, tergites 1V-V each with paired patches of 20-35 small circular
pores in two transverse rows on each side anterolaterally, pores with small conical collars internally,
tergites V-VI with intersegmental membrane invaginated into single-lobed pocket extending anteri-
orly 0.2—0.3 length of tergite V, tergite V three times as long as VI.

MALE TERMINALIA. — Ectoprocts each ovoid to crescent-shaped in dorsal view, apex rounded,
sclerotized ventrad of ventromedial lobe, ventromedial lobe slightly flattened dorsoventrally,
abruptly truncate along inner margin, covered with 20—30 short thickened setae; sternite [X pentago-
nal in ventral view, posterior margin broadly rounded medially; gonarcus with median lobe projecting
from dorsal surface, flattened anteroposteriorly, slightly curved anteriorly, twice as long as wide,
apex rounded; gonocoxites each three times as wide distally as basally in lateral view, abruptly wid-
ened at two-thirds length, apex truncate; mediuncus with anterior third transversely ovoid in lateral
view, 2—3 times as high as remainder, apex forked in posterior view; pseudopenal membrane ovoid in
posterior view, 1.3 times as long as wide, narrowed distally, covered with golden yellow dor-
sally-projecting microspines except for triangular region medially on basal third; hypomeres present
as paired small knobs; pseudopenis sclerotized, spinelike in posterior view, as long as pseudopenal
membrane and slightly longer than gonarcal median lobe, flattened anteroposteriorly.

FEMALE TERMINALIA. — Ectoprocts as long as gonocoxites; sternite VIII as long medially as lat-
erally, five times as wide as long in lateral view, slightly curved posteriorly, posterior margin truncate
medially; bursa with entrance entirely lightly sclerotized, remainder membranous to lightly
sclerotized; spermatheca with proximal section slightly wider than medial section, distal section 1.5
times as wide as proximal, diverticula present as short medially-directed lobe on inner margin, shorter
than width of spermatheca at that point, proximal section with first bend right laterad of medial sec-
tion, medial section with two coils, bent anteriorly before distal section, distal section with 1.5 coils,
fertilization canal duct sinuous; fertilization canal sloping basad of apex.

GEOGRAPHICAL DISTRIBUTION. — Costa Rica (LIMON); Panama (Canal Zone); French Guiana
(Saint Laurent du Maron).

ALTITUDINAL DISTRIBUTION. — Unknown for Costa Rica.

ADULT FLIGHT PERIOD. — March in Costa Rica; elsewhere in June, July, and December.

ETYMOLOGY. — Name derived from the Greek noun nymphe, which is a maiden-spirit inhabit-
ing trees, mountains, and bodies of water. The name is in reference to the forested regions of Central
and South America inhabited by this species.

TYPE MATERIAL. — Holotype male (USNM), four paratype males and four paratype females
(CHAH, CUAC, LSUC). Holotype label data: “COSTA RICA: Limon / Rio Uatsi, 8 km (air) / W
Bribri / 9.62°N, 82.90°W / 25.iii.1987, el. 60 m/ Holzenthal, Hamilton, Heyn,” “HOLOTYPE male /
Leptomantispa / nymphe Hoffman / det: K. M. Hoffman 1991.” Condition of holotype: color some-
what discolored, both eyes damaged, wings unspread, forewing length 10.3 mm. Paratypes have the
following data associated with them: COSTA RICA: Limon Prov., Rio Uatsi, 8 km (air) W Bribri,
9.62°N, 82.90°W, el. 60 m, 25-ITI-1987, Holzenthal, Hamilton, Heyn, 2 males and 3 females
(CUAC). PANAMA: Canal Zone, 6 km S Gattin, 30-XII-1972, Strauch, 1 female (CUAC); Canal
Zone, Pipeline Road, 10 km NW Gamboa, 9°11'N, 79°46'W, 25-VII-1977, H. A. Hespenheide, 1
male (CHAH); FRENCH GUIANA: Saint Laurent du Maroni Dept., 13 km SSE St. Laurent,
8-VI-1986, collected at mercury vapor light, E. G. Riley and D. A. Rider, | male (LSUC).

270 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES
Volume 53, No. 12

Leptomantispa pulchella (Banks, 1912) [new combination]
(Figs. 579, 585, 586, 608)

DIAGNOSIS. — Separated from other Costa Rican mantispines by the combination of prominent
setae over the entire length of the pronotum in lateral view with most setae arising flush with the
pronotal surface, the mesoscutellum brown medially and yellow laterally, the mesofemur with a lon-
gitudinal brown stripe ventrally, the forewing with subcostal space hyaline and the pterostigma light
red.

GEOGRAPHICAL DISTRIBUTION. — Southwestern Canada south to Costa Rica, including Cuba.

ALTITUDINAL DISTRIBUTION. — Unknown.

ADULT FLIGHT PERIOD. — May.

HABITAT PREFERENCE. — Unknown.

Xeromantispa Hoffman, new genus

TYPE SPECIES. — Mantispilla scabrosa Banks, by monotypy.

DESCRIPTION. — Head with shortest distance between eyes wider than width of one eye at ven-
tral margin of antennal sockets in frontal view, usually with distinct elliptical hump posteriad of
antennal bases medially, longer than wide and distinctly concave medially, antennal flagellum at
mid-length with flagellomeres twice as wide as long in anterior view. Pronotum straight in lateral
view, 4.4—5.0 times as long as wide at maculae in dorsal view, dorsal surface with numerous short
thick setae, setal bases not elevated. Pterothorax with mesoscutal furrow prominent, mesonotum with
numerous short thick setae, meso- and metascutelli with combined 3-5 pores on each side; legs each
with tarsomere I as long as tarsomeres II-V combined, tarsal claws with 5—6 teeth each; both wings
collectively on each side with combined 9-14 veins leaving RP posteriorly, forewing length
10.2-17.5 mm, sc-ra at 0.5—0.7 length of 3RA, Sc distal to sc-ra at 45—55° to RA distally, 3RA with
one c-ra crossvein distal to Sc, AA and 1 AP with posterior half asperous; hind wing with CuA strongly
angled toward AA, cu-aa crossvein either very short or absent and with CuA briefly fused with AA.
Male abdomen with tergite III lacking pores, tergites [V-V with 42—69 pores anterolaterally on each
side, most in transverse U-shaped row, anterior row longer than posterior row and nearly reaching
midline, several pores inside U and usually 1—2 isolated pores at mid-length laterally, pores circular
with short sclerotized conical collars internally, most pores in anterior row each with 1—3 long setae
arising from anterior margin which overlap pores in posterior row, other pores each with one seta aris-
ing from within; intersegmental membrane between tergites V-VI invaginated into single-lobed
pocket extending 0.4 length of tergite V, tergite V twice as long as VI. Male terminalia with each
ectoproct oval in dorsal view, apex broadly rounded ventromedial lobe rounded, wider than long, di-
rected medially, lightly sclerotized, with 35—46 short thickened setae; sternite [X subquadrate in ven-
tral view, wider than long, posterior margin produced into apically-concave subquadrate lobe with
posterolateral corners rounded, wider than long; gonarcus with medial lobe 1.5 times as long as wide,
somewhat square in dorsal view, flattened anteriorly, posterior surface flattened basally and anteri-
orly curved distally, apex rounded in posterior view, gonarcus with anterior margin produced dorsally
behind lobe into anteroposteriorly-flatttened flange, flange wider than high, rounded dorsolaterally
and deeply concave medially; gonarcal membrane unmodified; gonocoxites each as wide distally as
basally, apical fourth narrowed to a blunt spine dorsally and slightly bent laterally; mediuncus with
anterior two-fifths horizontally ovoid in lateral view, twice as high as distal three-fifths, apex forked
in posterior view; pseudopenal membrane ovoid in posterior view, sclerotized; hypomeres present as
paired very small granules, 1—2 on each side; pseudopenis triangular in posterior view, somewhat
shorter than pseudopenal membrane, twice as long as wide at base, lightly sclerotized, flattened
anteroposteriorly, base nearly as wide as mediuncal fork, apex pointed; hypandrium internum trian-

PENNY: LACEWINGS OF COSTA RICA 271

gular in ventral view, shallow and nearly straight in lateral view. Female terminalia with ectoprocts
1.2—1.5 times as long as gonocoxites; sternite VII entire, slightly longer medially than laterally, pos-
terior margin shallowly concave medially; bursa membranous; spermatheca with proximal section
slightly wider than medial section, distal section slightly narrower than proximal section, diverticula
present as low, narrow, transverse ridge, medial portion with from two bends to two complete coils
with connecting bend inside coils, fertilization canal duct curved basally, basal half dark, fertilization
canal sloping basad of apex.

CHROMOSOME COMPLEMENT. — Unrecorded.

IMMATURE STAGES. — Undescribed.

BiloLoGy. — Adults have been collected at lights and in pine-oak forests. Nothing is known of
immature stages.

GEOGRAPHICAL DISTRIBUTION. — Southwestern Unted States and northern Mexico.

ETYMOLOGY. — A combination of the Greek adjective xero, meaning arid, and mantispa as
coined by Illiger (1798), which is a combination of the Greek noun mantis, meaning soothsayer or
prophet, and the suffix pa, which is derived either from the Greek para, meaning near or beside, or
from the Latin par, meaning equal. The name refers to the arid climate prevalent in the region where
the type species occurs. Gender is feminine.

REMARKS. — Although this genus has a distribution outside the scope of this monograph, it was
deemed important to include the genus for completeness of coverage of the New World fauna. Costa
Rica contains all newly recognized New World genera, except Xeromantispa, and with its inclusion
the reader will have access to information covering all New World genera previously placed in the Old
World genus Mantispa.

Zeugomantispa Hoffman, new genus

TYPE SPECIES. — Mantis minuta Fabricius, 1775 by present designation.

DESCRIPTION. — Head with shortest distance between eyes narrower than width of one eye at
ventral margin of antennal sockets in frontal view, antennal flagellum at mid-length with
flagellomeres 1.3—1.8 times as wide as long in anterior view. Pronotum straight in lateral view,
5.27.2 times as long as wide at maculae in dorsal view, dorsal surface with numerous fine setae (Fig.
609), setal bases distinctly elevated. Pterothorax with mesoscutal furrow prominent, pterothoracic
scutella with 2—9 pores total on each side; legs each with tarsomere I distinctly longer than tarsomeres
II-V combined, pretarsal claws with 3—5 teeth each; both wings collectively on each side with com-
bined 6-17 veins leaving RP posteriorly, forewing length 6.0—16.5 mm, pterostigma with sc-ra at
0.5—0.7 length of 3RA (Fig. 590), distal end of Sc at 40—50° angle to RA, 3RA with one c-ra crossvein
distal to Sc, CuP with ends of terminal fork convergent and distinctly closer together than at base of
fork, AA and AP, simple, 1 AP asperous along AP»; hind wing with CuA strongly angled toward AA,
cu-aa crossvein very short or absent with CuA briefly fused with AA, jugal lobe with margin angulate.
Male abdomen with intersegmental membrane between tergites III-IV and IV-V invaginated into
paired bilobed pockets extending anteriorly 0.1—0.5 length of tergite V, each lobe with 1-10 pores.
Male terminalia with each ectoproct ovoid to crescent-shaped in dorsal view, apex rounded,
ventromedial lobe absent or barely present, ovoid patch of 20-47 short thickened setae
ventromedially, lightly sclerotized ventrad of setae; sternite [X pentagonal in ventral view, posterior
margin produced into short, slightly ventrally-directed, rounded lobe medially; gonarcus with dorsal
apex produced into medial lobe in lateral view; each gonocoxite up to three times wider distally than
basally in lateral view, apex narrowly rounded; mediuncus uniform in height in lateral view, apex
truncate or forked in posterior view; pseudopenal membrane triangular, unsclerotized; hypomeres
present as 2-4 small granules in longitudinal line on each side, often coalesced; pseudopenis
sclerotized, spinelike in posterior view, as long as pseudopenal membrane and median gonarcal lobe;

DI2 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES
Volume 53, No. 12

hypandrium internum triangular in ventral view, shallow and nearly straight in lateral view. Female
terminalia with sternite VII lengthened beneath tergite VIII; ectoprocts as long as or slightly shorter
than gonocoxites; sternite VII reduced, very short, medial third often present only as transverse line
of setae; bursa lightly sclerotized to membranous; spermatheca with proximal and medial sections
equal in width, distal section twice as wide as proximal, diverticulae present as transverse rod con-
nected medially to spermatheca, lateral arm sometimes reduced or absent, proximal section with first
bend laterad of medial section, medial section spiralling anteriorly for up to 1.5 loops before abruptly
bending posteriorly, distal section spiralling posteriorly for 1.0—1.5 loops and intertwining with me-
dial section, fertilization canal duct curved to sinuous, fertilization canal truncate or sloping basad of
apex.

CHROMOSOME COMPLEMENT. — Diploid chromosome complement of 20, consisting of 18
autosomes and 2 sex chromosomes based on Z. minuta (Hughes-Schrader, 1969). Sex chromosomes
of males are XY and those of females are XX.

IMMATURE STAGES. — All immature stages of Z. minuta have been described (Hoffman and
Brushwein 1992). The stalked eggs are oval with a caplike micropyle and are typical for mantispines.
First instars are diagnosed as follows: head capsule pale white, thoracic and abdominal terga with
paired reddish brown stripes laterally; mandible with one tooth on inner margin at three-fourths
length, antennal scape cylindrical, longer than wide; legs with tarsus longer than wide, unguitractor
process one-fourth as wide as tarsus and as long as tarsal claw, empodium twice as wide distally as ba-
sally; abdominal segments I-VI with setae D4, SD3, and L2 at most barely reaching posterior margins
of segments. Second and third instars are diagnosed as follows: legs with tarsus conical, claw present;
second instar with abdominal segment X longer than [X, tapered posteriorly; third instar with abdomi-
nal segment X as long as or longer than [X, spigot three times as long as base. Pupae are diagnosed as
follows: abdominal tergites III-IV each with setae 2—3 times as long as height of dorsal hooks in lateral
view, tergite IX with setae in two transverse rows laterally.

BIOLOGY. — Adults have been captured at lights and on various plants. Immatures of Z. minuta
have been recorded from the egg sacs of 29 spider species belonging to 14 families, including several
web-building species (Hoffman and Brushwein 1992); Z. viridula has been reared from the egg sacs
of two species in two families; and an adult of Z. compellens emerged from the egg sac of an unidenti-
fied orb weaver (Araneae: Araneidae). First instars of Z. minuta are obligate egg sac penetrators, are
very active, and have been shown to orient toward isolated spider egg sacs from a distance (Redborg
and MacLeod 1985). Valerio (1971) reported on some aspects of the biology of Z. minuta (cited as
Mantispa viridis Walker) in Costa Rica.

GEOGRAPHICAL DISTRIBUTION. — Eastern United States south to Argentina, including Cuba.

ETYMOLOGY. — A combination of the Greek adjective zeugo, meaning twinned, and mantispa
as coined by Illiger (1798), which is a combination of the Greek noun mantis, meaning a prophet or
soothsayer, and the suffix pa, which is derived either from the Greek para, meaning near or beside, or
the Latin par, meaning equal. The name is in reference to the two sets of invaginated intertergal mem-
branes between tergites III-[V and IV-V of males. Gender is feminine.

DISCUSSION. — This genus contains three species occurring from the eastern United States south
to Argentina, including Cuba. All three species occur in Costa Rica, and they can be distinguished by
the combination of prominent setae arising from distinct bumps on the pronotum and the forewing
with Sc distal to sc-ra at a 30° angle to RA

KEY TO COSTA RICAN ZEUGOMANTISPA

1. | Most wing veins uniformly brown, thoracic nota mostly brown, profemur usually with brown spot at base of major
Spihevanterolateralliys cms Sls Shes, us ec stec cssbtSiacieoon ss, stewed odes yh conten kl acs aa ee Rac Z. compellens

|’. Wing veins yellow or green, mottled with brown at vein junctions, thoracic nota yellow to green, profemur some-
times with red spots at base of major spine both antero- and posterolaterally ..................---. 2

PENNY: LACEWINGS OF COSTA RICA 213

2(1'). Antennal scape entirely green, forewing with aa-ap crossvein curved posteriorly (Fig. 590), nearly parallel with pos-
terior wing margin, | AA elliptical with posterodistal angle pointed, posterobasal angle distinctly different from

PISPCIOGIS ARO LC Mem eae rats nok gee ogni a TAs Sr Oe ieee ee eae a Z. minuta
2'. Antennal scape red posteriorly, forewing with aa-ap straight (Fig. 591), at distinct angle to posterior wing margin,
1AA rectangular, posterobasal angle similar to posterodistal angle ..................... Z. viridula

Zeugomantispa compellens (Walker, 1860) [new combination]
(Figs. 589, 594, 599)

DIAGNOSIS. — Separated from other Costa Rican mantispines by the combination of prominent
setae arising from distinct bumps on the pronotum, the mostly brown thoracic nota, wing veins mostly
uniformly brown, and the forewing with Sc distal to sc-ra at a 30° angle to RA.

GEOGRAPHICAL DISTRIBUTION. — Central Mexico south to southern Brazil. Within Costa Rica
specific localities are as follows: CARTAGO: Turrialba; GUANACASTE: Rincon National Park;
Santa Rosa National Park; 3 km SE Rio Naranjo; HEREDIA: La Selva Biological Station, 3 km S
Puerto Viejo, 10°26’N, 84°01'W; Finca La Selva Verde, 12 kmS Puerto Viejo; LIMON: Rio Banajo,
16 km WSW Bomba, 9.888°N, 83.167°W; Hacienda Tapezco, 29 air km W Tortuguero, 10°30'N,
83°47'W; Rio Uatsi, 8 km (air) W Bribri, 9.62°N, 82.90°W; PUNTARENAS: Golfito.

ALTITUDINAL DISTRIBUTION. — 40-300 m.

ADULT FLIGHT PERIOD. — February to April, June to September, November, and December.

HABITAT PREFERENCE. — Unknown.

Zeugomantispa minuta (Fabricius, 1775) [mew combination]
(Figs. 590, 595, 600, 609)

DIAGNOSIS. — Separated from other Costa Rican mantispines by the unique structure of the
forewing with the aa-ap crossvein curved posteriorly and nearly parallel with posterior wing margin
so that cell 1AA is elliptical with the posterodistal angle pointed and distinctly different from the
posterobasal angle. Other characters include: the combination of the antennal scape being entirely
green, prominent setae arising from distinct bumps on the pronotum, the yellow to green thoracic
nota, the wing veins yellow or green and mottled with brown at vein junctions, and the forewing with
Sc distal to sc-ra at a 30° angle to RA.

GEOGRAPHICAL DISTRIBUTION. — Eastern U.S.A. south to Venezuela, including Cuba. Within
Costa Rica it is known from the provinces of Cartago, Guanacaste, Heredia, Puntarenas, and San Jose.
Specific localities are as follows: CARTAGO: Cartago; 14.5 km (9 mi) NW Esparta, Turriabla, ICA
Station; Turrialba; GUANACASTE: near Arenal, 10°20'N, 84°50'W; Playas del Coco; Taboga; Ha-
cienda Tempisquito (Pelon de la Altura) | km NE of km 265, rt 1, 10.84°N, 85.56°W; Las Canias, Rio
Corobici; S. Cafias, Experiment Station; La Pacifica, nr. Cafias; 2 mi NW of Cafias; Hacienda La
Pacifica, 5 km NW Cafias; Hacienda La Pacifica, 6 km N Cafias; 14 km S Canas; 20 km SW Canas;
La Pacifica, 37 km(23 mi) NW Cajias; 3 km SE Rio Naranjo; Parque Nacional Guanacaste, El Hacha,
Quebrada Alcornoque, 11.009°N, 85.577°W; Parque Nacional Guanacaste, Maritza, Rio
Tempisquito, 10.958°N, 85.497°W; Parque Nacional Rincon de la Vieja, Rio Negro, 10.765°N,
85.313°W; Rio Gongora (sulfur mine), 4 km (air) NE Quebrada Grande, 10.887°N, 85.470°W; Playa
Hermosa; Santa Rosa National Park; La Pacifica, 4 km NW Cafias; HEREDIA; San Rafael Vara
Blanca; PUNTARENAS: Boca de Barranca; Monteverde; 8.1 km (5 mi) E of Judas, CR-38; 25.5 km
(15.8 mi) W of San José, 8.1 km (5 m1) E Judas, CR-38; SAN JOSE: San José, Universidad de Costa
Rica; San José; Fadra La Caja, 8 km W San José.

ALTITUDINAL DISTRIBUTION. — 100-2200 m.

ADULT FLIGHT PERIOD. — Year round in Costa Rica.

HABITAT PREFERENCE. — Unknown.

274 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES
Volume 53, No. 12

REMARKS — Three names recorded in the literature from Costa Rica are here synonymized with
Zeugomantispa minuta. They are: Mantispilla flavicornis Navas, 1930 [new synonymy]; Mantispilla
rubricata (Navas, 1924) [new synonymy], and Mantispilla viridata (Navas, 1924) [new synonymy].
The North American name Mantispa viridis Walker, 1853 is also a new synonym of Zeugomantispa
minute.

Zeugomantispa viridula (Erichson, 1839) [new combination]
(Figs. 541, 591, 596, 601)

DIAGNOSIS. — Separated from other Costa Rican mantispines by the combination of the
antennal scapes being red posteriorly, prominent setae arising from distinct bumps on the pronotum,
the yellow to green thoracic nota, wing veins yellow or green and mottled with brown at vein junc-
tions, the forewing with Sc distal to sc-ra at a 30° angle to RA, and the forewing with aa-ap straight and
at a distinct angle to posterior wing margin so that cell 1 AA is rectangular with the posterobasal angle
similar to posterodistal angle.

GEOGRAPHICAL DISTRIBUTION. — This species occurs from central Mexico south to Argentina.
Within Costa Rica specific localities are as follows: ALAJUELA, 2 km NE Biagua; Parque Nacional
Rincon de la Vieja, Quebrada Provision, 10.769°N, 85.281°W; Reserva Forestal San Ramon, Rio San
Lorencito and tribs., 10.216°N, 84.607°W; GUANACASTE: Comelco Property; Las Cafias, Rio
Corobici; La Pacifica, nr. Cahas; Hacienda La Pacifica, 5 km NW Canas; 14 km S Canas; 3 km SE
Rio Naranjo; Santa Rosa National Park; Quebrada Garcia, 10.6 km ENE Quebrada Grande,
10.862°N, 85.428°W; HEREDIA: Finca La Selva Verde, 12 km S Puerto Viejo; La Selva Biological
Station, 3 km S of Puerto Viejo, 10°26'N, 84°01'W; 1 km S Puerto Viejo; Rio Bijagual on road to
Magsasay, 10.408°N, 84.076°W; San Rafael Vara Blanca, 2200 m; LIMON: Pandora; Rio Banaiio,
16km WSW Bomba, 9.888°N, 83.167°W; Rio Uatsi, ca. 8 km (air) W Bribri, 9.62°N, 82.90°W;
PUNTARENAS: 3.2 km (2 m1) SW San Vito de Java; La Amistad National Park, 8°57'N, 82°50'W;;
Finca Las Cruces, 6 km S San Vito de Java.

ALTITUDINAL DISTRIBUTION. — 60-2200 m.

ADULT FLIGHT PERIOD. — Every month except August.

HABITAT PREFERENCE. — Unknown.

SUBFAMILY SYMPHRASINAE

Two of the three genera of the Symphrasinae occur in Costa Rica, each represented by a single
species. Larvae have three instars and are hypermetamorphic with the first instar campodeiform and
the last two scarabaeiform (Parker and Stange 1965). Members of the Symphrasinae have been reared
from the nests of aculeate Hymenoptera and from pupae of Lepidoptera, and have been associated
with the pupae of Coleoptera and Diptera.

Plega Navas, 1927

Plega contains 14 species occurring from the southwestern United States south to Bolivia and
Brazil, only one of which is known from Costa Rica. Adults of the genus have been associated with
pupae of Scarabaeidae (Coleoptera), Asilidae (Diptera), Noctuidae (Lepidoptera), and various
Hymenoptera (Anthophoridae, Megachilidae, Sphecidae, and Vespidae: Eumeninae) (Parker and
Stange 1965; Werner and Butler 1965; Penny 1982).

PENNY: LACEWINGS OF COSTA RICA 2S

Plega hagenella (Westwood, 1867)
(Figs. 538, 543, 548, 553)

DIAGNOSIS. — Separated from other Costa Rican mantispids by the combination of a sub-basal
spine on the profemur and its two-piece pronotum.

GEOGRAPHICAL DISTRIBUTION. — This species is recorded from Costa Rica, Colombia and
Brazil. Within Costa Rica it was recorded by Navas (1924) as Symphrasis signata Hagen from an un-
specified locality, and has been subsequently collected in the specific localities as follows:
GUANACASTE: S Canfas; 14 km S Canfas; 3 km SE Rio Naranjo; Santa Rosa National Park;
HEREDIA: La Selva Biological Station, 3 km S Puerto Viejo, 10°26'N, 84°01'W.

ALTITUDINAL DISTRIBUTION. — 180-300 m.

ADULT FLIGHT PERIOD. — January to March, May to July, and November to December.

HABITAT PREFERENCE. — Unknown.

Trichoscelia Westwood, 1852

Trichoscelia contains 13 species occurring from southern Mexico south to Argentina, only one of
which is known from Costa Rica. Penny (1982) revised the Brazilian species, but the Central Ameri-
can and northern South American species are poorly known and it is not possible to confidently iden-
tify species at this time. Three species have been described from southern Mexico and one from
northern Colombia. I have chosen to assign the present species from Costa Rica to the oldest available
name, which is the one from Colombia. Future revisions of Trichoscelia may or may not support this
decision. Adults of this genus have been reared from the nests of Polybia species (Hymenoptera:
Vespidae: Polistinae) (Penny 1982).

Trichoscelia remipes (Gerstaecker, 1888)
(Figs. 537, 539, 542, 549, 554)

DIAGNOSIS. — Separated from other Costa Rican mantispids by the lack of a sub-basal spine on
the profemur.

GEOGRAPHICAL DISTRIBUTION. — This species was described from Colombia and is now
known to occur in Costa Rica as well. Within Costa Rica specific localities are as follows:
GUANACASTE: Parque Nacional Rincon de la Vieja, Rio Negro, 10.765°N, 85.313°W;
HEREDIA: La Selva Biological Station, 3 km S Puerto Viejo, 10°26'N, 84°01'W.

ALTITUDINAL DISTRIBUTION. — 810 m.

ADULT FLIGHT PERIOD. — March, May, and July.

HABITAT PREFERENCE. — Unknown.

FAMILY MYRMELEONTIDAE

by
Lionel A. Stange

The members of this family are called antlions in reference to the characteristics of the larvae.
The actual name is derived from ancient references to animals with the head ofa lion and the body of
an ant, but today we think of antlions as being voracious predators on ants. Antlion larvae are superbly
suited to digging in the soil or loose sand and nearly all species, where the biology is known, have this
digging ability. However, most people associate antlions with the funnel shaped pitfall traps that can
be abundant in areas such as sand dunes. Many hundreds of years ago the Mimbres Indians of New

276 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES
Volume 53, No. 12

Mexico depicted antlion larvae on their pottery. Most of the pitfall builders belong to the tribe
Myrmeleontini (but some others are found in other tribes, although they often have different struc-
tures to their pitfalls which in the case of certain Brachynemurini are built in hardpan soils. In Costa
Rica there are four tribes of antlions, but most species belong to the tribe Nemoleontini, which do not
make pitfall traps. The larvae of this group live in sand (Purenleon, Dimarella), in cave mouths
(Eremoleon) or in sawdust in rotten trees (Glenurus). The tribe Acanthaclisini is represented by the
genus Vella, which are giant antlions. Their larvae live in deep sand and can only move backwards and
often prey on the larvae of Myrmeleon which only move backwards as well. The fourth tribe of ant-
lions represented in Costa Rica is the Brachynemurini which are represented by two genera, Dejuna
and Abatoleon. The larvae of these genera are free living in sand. Adult antlions fly at night and appar-
ently copulate and oviposit in the early evening and then spend the rest of the night looking for food
which may consist of sessile insects such as aphids, or lepidopterous caterpillars, or even other antlion
adults. Some species are known to be attracted to night blooming flowers and may eat pollen, but very
little is known of the Costa Rican species. During the day adults rest on grasses (Dejuna, Abatoleon,
Myrmeleon), on rock surfaces or on tree trunks or posts (Vella). Many species of antlions are attracted
to lights but others are not . The rarer species can most easily be collected by finding the larvae. Larvae
can be sifted from the sand or exposed in restricted habitats such as tree bases or in cave mouths by
blowing away the substrate with an aspirator. Usually the larvae can be reared easily.

KEY TO THE GENERA OF COSTA RICAN MYRMELEONTIDAE

ADULTS

1. | Metafemur with elongate sense hair (Fig. 616); hind wing vein CuA unites with posterior fork of MP, shortly after
fork (Fig. 613); distal segment of labial palpus with sensory area (palpimacula) elongate and slit-like (Fig. 614); large
(forewing length over 50 mm), robust and hairy insects (Fig. 615). . . . (Tribe Acanthaclisinl) ........ Vella

1’. Metafemur without elongate sense hair; hind wing vein CuA running to hind margin, usually connected to posterior
fork of MP, by crossveins; distal segment of labial palpus with sensory area oval-shaped; small to medium species
(forewing less than 50 mm), usually fragile and not extremely hairy .................2..5---- 2

2(1'). Forewing vein 2A evenly curved from base to hind margin; forewing vein CuP originates well distad of basal

crossvein (Fig. 611); male ectoproct with distinctive postventral lobe. . . (Tribe Brachynemurini)........ . 3)
2’. Forewing vein 2A runs close to vein 1A for a short distance, then bends at sharp angle toward 3A; forewing vein CuP
originates at level of basal crossvein (Fig. 612); male ectoproct without postventral lobe (except Dimarella) .... 4

3(2). Ocular rim with 1 or more setae that project over eye; forefemur with row of distally swollen setae on exterior face;
male ectoproct with postventral lobe less than three times longer than middle diameter; female abdomen longer than
HOTE WINE. 595. 2S SI ba Gees ees a See ee eS Se oh SS a Abatoleon
3’. Ocular rim without setae; forefemur with or without row of setae, if present, setae not distally swollen; male
ectoproct with postventral lobe more than eight times longer than middle diameter; female abdomen shorter than
FORE WANES Meee es Ae. Te SE a, Te SP ec ec Dejuna

4(2'). Hind wing with radial sector arising beyond cubital fork, four or more presectoral crossveins present; male pilula
axillaris present; distal tarsomere with ventral setae less than one-half as long as tarsomere diameter... ..... -
(nibeiMyrmeleontini\in= % So 6 Ge oe aco eee oe © hee Oe ey Oe ee Myrmeleon
4’. Hind wing with radial sector arising before cubital fork, only one presectoral crossvein present; male pilula axillaris
absent; distal tarsomere with ventral setae at least as long as tarsomere diameter. . . (Tribe Nemoleontini) ... . 5

5(4’). Antennal socket separated by more than greatest width of antennal pedicel from ocular rim; posterior fork of
forewing vein CuA and vein CuP + 1A parallel with each other and hind margin for long distance; male ectoproct
Hailin Ponsa oor 5 a5 nels a6 His oo Obl HSS Slo SiblaS Bo DB aS a5 6B os oe Dimarella

5’. Antennal socket separated by less than greatest width of antennal pedicel from ocular rim; posterior fork of forewing
vein CuA at an oblique angle to hind margin; male ectoproct without elongate postventral lobe... .......-.- 6

PENNY: LACEWINGS OF COSTA RICA 277

6(5’). Forefemur swollen, at widest point much wider than interantennal distance with abundant pubescence, especially on

closing surface; femoral sense hair of foreleg over one-half lengthoffemur ................ Purenleon
6’. Forefemur slender and elonsgate, at widest point about equal at most to interantennal distance, with sparser pubes-
cence; femoral sense hair of foreleg less than one-third length offemur.................-....... 7

7(6'). Pretarsal claws capable of closing against distal tarsomere, which has brush of specialized setae on ventral surface on
Sie MONG 11a amen th eee EAS 62 Pes eae. os oe AS, gS hoes See Ee aA oe BVP Sule te 2 et Elachyleon

Weeenrctarsal claws not capable of closing against distalitarsomere -..---.-..--:-+ 2-2 see eee eee se 8
8(7'). Forewing costal area with single series of cells for most of distance; apical one-fifth of hind wing and usually
forewing with predominant dark brown suffusion in marked contrast to basal one-fifth (Figs. 629-631) . . Glenurus
8’. Forewing costal area with double series of cells for most of distance (Figs. 633, 636); apical one-fifth of wings pre-
dominantly transparent, sometimes with many scattered dark brown spots .............-... Eremoleon
LARVAE

(unknown: Elachyleon punctipennis; Dimarella garciai,; Eremoleon species)

1. Labial palpus shorter than basal width of mandible; mesothoracic spiracle not borne on a tubercle; head without
Holichusterssicamonly-moyverbackwardSs 2). cou. Gil a) clean Geos 2). elec Bete tee conn Ale eee See aie 2

1’. Labial palpus longer than basal width of mandible or mesothoracic spiracle borne on a tubercle; head often with

dolchasterssmove tonwards and backwards: 94) 68%. Geb) 8 cele nee os Ee alg ae Sen, ee eee 3
2(1). Mandible with some setae on outer lateral margin as long as or longer than greatest mandibular width; sternite VIII
with pair of inconspicuous submedian processes near posterior margin; make pitfall traps... . 2... . Myrmeleon
2'. Mandible with longest setae on outer lateral margin less than one-half greatest mandibular width; sternite VIII
without tecthon subapicalimarcin; donot make pitfall traps! - . . . . . 2. we ee: Vella fallax
3(1'). Mandible with distal tooth shorter than middle tooth and usually set at different angle than other teeth... .....
((Riaclnanerniinion) se lool ol cals dhe elon eon NS CAPE CeRE! Gach Cp Some emnOnY Pon MEE ONC anes Sheses aye torrns ty cha c 4
3’. Mandible with distal tooth as long as or longer than middle, teeth parallel... . . (Nemoleonimni) See eee 5
4(3). Sternite VIII with submedian pair of sclerotized processes near posterior margin. ............. Abatoleon
PeESLCUINLC AV wth out SGlerotizedsprocesseS) = = iss sas 0s : S02 ole, 2) © lcs) eee) en) ee Ee Dejuna
Ten mivanciblenmthitwotceth= abdomenswithiscOll 3 20.05 < se ae 8 ee eee eee Glenurus
3 Manuiblenwithithnee tecthabdomenkswithourSCOll. 454-04 © alee se @ stele: © oie cy ie ye eee ene 6
Gon) slvateral abdominal spiracles situated on wellraised tubercles 7 2... J 2 2 eee ee ii
6. Lateral abdominal spiracles more or less sessile, not on tubercles ..........-.-++++-+-++++-+--- 8

7(6). Lateral abdominal spiracles with papilliform tubercles beset with short dolichasters .........-...-.-.--.
PU 6 ee Ty ase rn i t Pee ee MR eh oi sh op Bee Purenleon debilis, Purenleon iniquus

7’. Lateral abdominal spiracles with bulbous tubercles not beset with dolichasters .......... Dimarella angusta
Sone lawvacliveunderrockOvethangsiOnin GAaVeESis, | 5 65) 2 2)5 20). 2) es ee ee ee ee Eremoleon
8’. Larvae do not live under rock overhangs nor cave mouths ......... Dimarella mixteca, Purenleon parallelus

TRIBE ACANTHACLISINI

This tribe is composed of 14 described genera found in most of the temperate and tropical world.
Most of the species are large, robust and hairy, with well-developed tibial spurs and pretarsal claws.
This is true of the only genus of the tribe in Costa Rica, Vella Navas.

978 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES
Volume 53, No. 12

Vella Navas, 1913

This genus contains six species restricted to the Western Hemisphere. Vella fallax (Rambur) ex-
tends from southwestern United States to Argentina and is the only species recorded from Costa Rica.
The adults are attracted to lights and during the day rest on tree trunks or on posts. The larvae live in
deep sand and can only move backwards in search of their prey, which may include other antlion lar-
vae.

Vella fallax (Rambur, 1842)
(Figs. 613-616)

DIAGNOsIS. — Large antlions with a body length between 45 and 60 mm and a wing span reach-
ing 160 mm. The wings are relatively unmarked with a biareolate forewing costal area (Fig. 613). One
unique venational character of this species in Costa Rica is the hind wing vein CuA which unites with
the posterior fork of MP) shortly after fork. Another unique feature of this species (in Costa Rica) is
the presence of an elongate femoral sense hair on the hind femur (Fig. 616).

GEOGRAPHICAL DISTRIBUTION. — U.S.A. (AZ, TX) south to Argentina. In Costa Rica there are
records from GUANACASTE: Santa Rosa National Park.

ALTITUDINAL DISTRIBUTION. — 300 m (Guanacaste).

ADULT FLIGHT PERIOD. — July to January in Costa Rica.

HABITAT PREFERENCE. — This species should be found in many lowland areas where there ex-
ists enough deep sand to support the larvae. It should be found in some of the beach areas of Costa
Rica.

TRIBE BRACHYNEMURINI

This tribe is restricted to the Western Hemisphere. Only one genus (Dejuna Navas) has been col-
lected in Costa Rica to date, but it is likely that species of Abatoleon Banks will be found in the future,
since there are records from Mexico and Colombia. Members of this tribe differ from all other groups
in Costa Rica in having the forewing vein CuP originating well distad of basal crossvein (sometimes
obsolete) (see Fig. 611). Also, the male ectoproct has an elongate postventral lobe (found only in
Dimarella in Costa Rica) and the forewing vein 2A runs ina fairly even curve to hind margin. The lar-
vae have a distinct mandibular dentition, in which the distal tooth is shorter than the middle tooth and
is usually set at a different angle than the other teeth.

Dejuna Navas, 1924

The species of this genus are known only from the arid tropical areas of Mexico and Central
America. The adults are mostly straw colored and rest on dry grasses during the day except for the ab-
errant species, Dejuna mimica, which is brown and has been found resting on dry plant stems. In Costa
Rica, the males are easily distinguished from other species (except those of Dimarella) by the
ectoproct which has an elongate postventral lobe that is more than four times longer than its middle di-
ameter. The males of Dejuna have a well-developed pilula axillaris which is absent in Dimarella.

KEY TO COSTA RICAN DEJUNA

1. | Forefemur with row of white bristles for most of distance on exterior face (Fig. 628); tibial spurs of midleg equal to at
least length of basal two tarsomeres; distal palpimere of labial palpus nearly as long as interocular distance below
antennae’ (FIGHG26)i 5 ices ots au © we. Gee arse sy sh get eee ee cr chal eRe) ete lc ot nent rae D. setosa

PENNY: LACEWINGS OF COSTA RICA 279

1’. Forefemur without bristles on exterior face except sometimes subapically (Fig. 624); tibial spurs of midleg no longer
than basitarsus: distal palpimere of labial palpus much shorter than interocular distance (Fig.623) ......... 2

2(1'). Brown colored (Fig. 625); femoral sense hair absent; setae on legs abbreviated, none longer than width of femur
RR aC | Mn ees re tt Geen ae ee BPR YT ele Deas, ee ee © oes D. mimica

2'. Straw colored; femoral sense hair present and well developed; some setae on legs longer than width of femur .. . .
arene: waren tars fet oge tte gms cite biel it aeoe ee Wn oe et SP eS Ee ay Bee Ie D. fenestrata

Dejuna fenestrata (Banks, 1913) [new combination]
(Fig. 620)

DIAGNOSIS. — This straw colored species is closest in appearance to Dejuna setosa but has a
shorter labial palpus, lacks the row of white bristles on the exterior face of the forefemur and has
broader wings (Fig. 620). The adult length ranges from about 19 mm (female) to 29 mm (male).

GEOGRAPHICAL DISTRIBUTION. — This species is known from Guatemala, Honduras and Costa
Rica. The type specimen was collected at ALAJUELA: Turricares; and other specimens were col-
lected in GUANACASTE: Santa Rosa National Park.

ALTITUDINAL DISTRIBUTION. — 300 m.

ADULT FLIGHT PERIOD. — October to January.

HABITAT PREFERENCE. — The adults rest on grasses during the day. The species congregate in
fairly large numbers.

Dejuna mimica (Stange, 1970) [new combination]
(Figs. 621, 623-625)

DIAGNOSIS. — This relatively small antlion (Fig. 621) (body length 19 to 23 mm) has reduced
leg setae and lacks the femoral sense hair (Fig. 624). This latter feature is unique among Costa Rican
antlions. The species resembles those of the genus Atricholeon Stange which is found in the south-
western United States and Mexico.

GEOGRAPHICAL DISTRIBUTION. — This species is known from southern Mexico, Guatemala,
Honduras, and Costa Rica. The species was collected in Costa Rica in GUANACASTE: Santa Rosa
National Park.

ALTITUDINAL DISTRIBUTION. — 300 m.

ADULT FLIGHT PERIOD. — March and April.

HABITAT PREFERENCE. — Unknown.

REMARKS. — Stange (1970) found males resting with the abdomen held out at about a 45 degree
angle from the plant stem.

Dejuna setosa (Stange, 1970) [new combination]
(Figs. 622, 626-628)

DIAGNOSIS. — One outstanding diagnostic character of this species is the elongate labial palpus
that is about equal in length to the interocular distance at the level of the antennae (Fig. 626). Also, the
forefemur has a row of white bristles (Fig. 628) and other bristles are found on the mesoscutellum and
posterolateral margin of the pronotum. This species (Fig. 622) is about the same size as D. fenestrata
(body length 25 to 29 mm) and is also straw colored (Fig. 627).

GEOGRAPHICAL DISTRIBUTION. — This species is known from southern Mexico, Nicaragua and
Costa Rica. The only locality known for this rather rare species in Costa Rica is GUANACASTE:
Santa Rosa National Park.

ALTITUDINAL DISTRIBUTION. — 300 m.

280 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES
Volume 53, No. 12

ADULT FLIGHT PERIOD. — March and April.
HABITAT PREFERENCE. — Unknown.

TRIBE MYRMELEONTINI

This is a very distinct and morphologically conservative tribe that has larvae that can build pitfall
traps in the sand and only move backwards. One unique structural characteristic of this tribe is the
chaetotaxy of the distal tarsomere. All setae on the ventral side of the distal tarsomere are very short,
less than one-half the diameter of the tarsomere. Also, the hind wing radial sector arises beyond the
cubital fork with at least four presectoral crossveins.

Myrmeleon Linnaeus, 1767

This cosmopolitan genus has 165 described species and thus rates as the largest genus in the fam-
ily. However, much taxonomic revision 1s needed for most areas of the world. Larvae from all parts of
the world appear to be structurally nearly identical, and the adults offer few striking characters, al-
though the male genitalia appear to be distinctive for each species. A vast literature exists concerning
the pit making activities of Myrmeleon, especially in Europe and the United States. Usually there are
several species at any one locality. The species (e.g., Mvrmeleon timidus) make pitfall traps, occasion-
ally in open areas, but most Costa Rican species make them in protected areas such as under rock over-
hangs.

KEY TO COSTA RICAN MYRMELEON

1. Forewing with costal area greatly expanded at pterostigma, (over twice as broad as above radial sector), and with
several crossveins basad to pterostigma interconnected (Fig. 617) (subgenus Neohornius Navas) . M.(N.) perspicuus

1’. Forewing with costal area not greatly expanded at stigma, (at most 1.5 times wider than above radial sector), and
without interconnected crossveins (rarely 1, 2) (Fig. 618, 619) (subgenus Myrmeleon L.) .............. 2

2(1). Moderate-sized species with body length ranging from 29 to 34 mm; face with broad black band below antennae

extending entrallyonlyato above levellotitentonialipitsees eee aie Gene, eee eee M. (M.) uniformis
2'. Body length 23 to 27 mm; face with broad black band below antennae extending ventrally to below level of tentorial
[SHG es <a trie Get ee nek Hoses B teres st eee a een ne eae PW IREE RNR Le oo M. (M.) timidus

Myrmeleon (Neohornius) perspicuus Gerstaecker, 1893
(Fig. 617)

DIAGNOsIS. — The broadened forewing costal area near the pterostigma with interconnected
crossveins (Fig. 617) distinguishes this species from other Myrmeleon in Costa Rica. The body size
ranges from about 30 to 35 mm.

GEOGRAPHICAL DISTRIBUTION. — Described from Panama (Chiriqui) and in Costa Rica known
from CARTAGO: Santa Teresita de Lajas; Turrialba; HEREDIA: La Selva Biological Station: SAN
JOSE: San Antonio-Dasimp; Santa Ana.

ALTITUDINAL DISTRIBUTION. — Santa Ana in the Central Valley and Turrialba are both at
mid-elevations. La Selva is in the lowlands to the north.

ADULT FLIGHT PERIOD. — July to October.

HABITAT PREFERENCE. — Unknown.

PENNY: LACEWINGS OF COSTA RICA 281

Myrmeleon timidus Gerstaecker, 1888
(Fig. 618)

DIAGNOSIS. — This is the smallest species of Myrmeleon in Costa Rica (Fig. 618) with speci-
mens not exceeding 27 mm in body length. This species belongs to the exclusively New World spe-
cies group of M. metuendus, characterized by the development of a sclerotized process ventrally on
the male tergite [X. In M. timidus this process is black and bilobed.

GEOGRAPHICAL DISTRIBUTION. — Mexico south to northern Chile on the western side of the An-
des Mountains. Also found on the Galapagos Islands. In Costa Rica, there are records from
GUANACASTE: Santa Rosa National Park; and PUNTARENAS: the beaches at Quepos. This spe-
cies is probably found throughout most of Costa Rica, wherever the proper habitat is found.

ALTITUDINAL DISTRIBUTION. — Sea level to at least 400 m.

ADULT FLIGHT PERIOD. — December to June.

HABITAT PREFERENCE. — The pitfall traps are usually made in protected areas, except in very
dry areas, where they may be made in the open.

Myrmeleon uniformis Navas, 1920
(Fig. 619)

DIAGNOSIS. — This species is larger (Fig. 619) (body length more than 28 mm) than M. timidus
(body length less than 27 mm) and tergite IX is not produced ventrally in the male. From M.
perspicuus, which is about the same size, the lack of interconnected costal crossveins before the
pterostigma provides a reliable recognition character.

GEOGRAPHICAL DISTRIBUTION. — Mexico south to Costa Rica. In Costa Rica there are records
from PUNTARENAS: Monteverde; and SAN JOSE: Escazu.

ALTITUDINAL DISTRIBUTION. — Monteverde is at about 1500 m elevation, whereas Escazu is a
suburb of San José at a somewhat lower elevation in the Central Valley.

ADULT FLIGHT PERIOD. — August.

HABITAT PREFERENCE. — This species constructs its pitfall traps in protected areas such as un-
der rock overhangs and at cave mouths.

REMARKS. — Although Myrmeleon mexicanus Banks was recorded from Costa Rica by Banks
(1914), the specimen probably is Myrmeleon uniformis Navas, which 1s closely related and perhaps a
synonym. Substantiated records of M. mexicanus are known only from north of the Neovolcanic Pla-
teau of Mexico, those of M. uniformis south of the Neovolcanic Plateau with no sympatry known. The
name Myrmeleon leptaleus Navas, 1924 that was previously recorded from Costa Rica is a new syn-
onym of Myrmeleon uniformis.

TRIBE NEMOLEONTINI

This is the largest tribe in the family both in numbers of genera and species. The wing venation
distinguishes this tribe from others found in Central America. The hind wing radial sector arises close
to the wing base and there is only one presectoral crossvein. The forewing vein 2A runs along 3A fora
short distance before angling abruptly toward the hind margin. This condition is also found in the
Myrmeleontini. There are five genera recorded from Costa Rica. Dimarella Banks is the most distinc-
tive because the male ectoproct has an elongate postventral lobe (absent in other genera of this tribe),
the antennal scrobe is widely separated from the eye, and the pretarsal claws are modified to close
against the ventral surface of the distal tarsomere (found also in Elachyleon, Fig. 648). Glenurus
Hagen contains species with considerable wing markings (Fig. 629-631). Its larvae are very distinc-
tive in having only two mandibular teeth. Eremoleon Banks contains two species in which the larvae

282 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES
Volume 53, No. 12

live in cave mouths or rock overhangs. The larvae of Purenleon Banks live in sand. The fifth genus,
Elachyleon Esben-Petersen, is unknown as to larval habitat and appears to be relatively rare.

Dimarella Banks, 1913

This genus was recently revised by Miller and Stange (1989) and contains three species found in
Costa Rica. The larvae live in sand and resemble those of Purenleon. The adults are very distinctive in
having the antennal bases far removed from the eye (by more than width of antennal scrobe), having
an elongate postventral lobe developed from the male ectoproct, and in having the pretarsal claws
modified to close upon the ventral surface of the distal tarsomere.

KEY TO COSTA RICAN DIMARELLA

1. Forewing vein CuP+1A reaching hind margin near level of origin of radial sector; forewing posterior area with one

series of cells and narrower than cubital area at level of radial sector (Fig.655) .............. D. mixteca
1’. Forewing vein CuP+1A reaching hind margin well beyond level of the origin of the radial sector; forewing posterior

area with two series of cells and wider than cubital area at origin of radial sector (Figs. 649,652) .......... 2,
2(1)sForetemurcylindricalyat least nme times longer thaniwide(Figs650)) @ 2) e)o eee eee D. angusta
a Forefemur greatly swollen, about five times longer than wide (Fig.653) .................. D. garciai

Dimarella angusta (Banks, 1908)
(Figs. 649-651)

DIAGNOSIS. — This species has body length between 20 to 25 mm and the many antennal
flagellomeres are longer than wide (Fig. 651). The forefemur is about nine times longer than wide,
which separates it from D. garciai which has similar broad wing venation. From D. mixteca, the wing
venation (Fig. 649) is distinctive.

GEOGRAPHICAL DISTRIBUTION. — Described from Ecuador, widespread in South America (Co-
lombia, Venezuela, Brazil, Peru, Bolivia) and known from Panama and Costa Rica. Records from
Costa Rica are from HEREDIA: La Selva Biological Station, 3 km S Puerto Viejo; LIMON: Rio
Reventazon; and PUNTARENAS: Boca de Barranca.

ALTITUDINAL DISTRIBUTION. — Mid-elevations.

ADULT FLIGHT PERIOD. — March to June.

HABITAT PREFERENCE. — Unknown.

Dimarella garciai (Navas, 1932)
(Figs. 652-654)

DIAGNOSIS. — The greatly swollen forefemur (Fig. 653), which is about five times longer than
greatest diameter, is distinctive among Costa Rican species. Also, the mesoscutum lacks the promi-
nent white bristles found in other species of Dimarella in Costa Rica. The size of this species is about
the same as D. angusta.

GEOGRAPHICAL DISTRIBUTION. — Known from Ecuador, Costa Rica, and Panama. The only lo-
cality for this species in Costa Rica is GUANACASTE: Santa Rosa National Park.

ALTITUDINAL DISTRIBUTION. — 300 m.

ADULT FLIGHT PERIOD. — December.

HABITAT PREFERENCE. — Unknown.

PENNY: LACEWINGS OF COSTA RICA 283

Dimarella mixteca Miller, 1989
(Figs. 655-657)

DIAGNOSIS. — The body length of this species averages a little smaller (21 mm) than other spe-
cies in Costa Rica. It belongs to a species group centered in Mexico and has distinctive wing venation
among Costa Rican species. The forewing vein CuP+1A is short and reaches the hind margin at about
the level of the origini of the radial sector. Also, the forewing posterior area is very narrow and con-
tains only one series of cells (Fig. 655).

GEOGRAPHICAL DISTRIBUTION. — Mexico, Honduras, and Costa Rica.

ALTITUDINAL DISTRIBUTION. — Unknown.

ADULT FLIGHT PERIOD. — Unknown.

HABITAT PREFERENCE. — Unknown.

Elachyleon Esben-Petersen, 1927

This is a poorly known genus probably closely related to Eremoleon. The larval habitat for spe-
cies of this genus has not been discovered. The species have elongate legs and the pretarsal claws are
modified to fold against the specialized setal patch on the ventral side of the distal tarsomere. This
condition is found elsewhere in Costa Rica only in the genus Dimarella. There are two described spe-
cies in the genus, the type species, Elachyleon punctipennis, which ranges from Mexico to Argentina,
and Elachyleon gentilis (Banks) known only from Guyana.

Elachyleon punctipennis Esben-Petersen, 1927
(Figs. 610, 646-648)

DIAGNOSIS. — This species has a bright yellow pronotum and is about 20 mm in body length.
The legs are very long (Fig. 648) and the pretarsal claws can fold against a specialized setal patch on
the ventral side of the distal tarsomere (Fig. 648). The setae in the setal patch are hooked apically (in
contrast to those found in Dimarella).

GEOGRAPHICAL DISTRIBUTION. — Mexico south to Argentina. The type was described from
Trinidad, but a more prominently marked subspecies (wings, Figs. 610, 646) was described from
LIMON: Las Mercedes, Costa Rica. Other records from Costa Rica are GUANACASTE: Parque
Nacional Guanacaste; and HEREDIA: 12 km S Puerto Viejo.

ALTITUDINAL DISTRIBUTION. — 500 m (Puerto Viejo).

ADULT FLIGHT PERIOD. — April, July, and September in Costa Rica.

HABITAT PREFERENCE. — Unknown.

REMARKS. — Elachyleon punctipennis pulchellus Esben-Petersen, 1932 is a new synonym of
Elachyleon punctipennis.

Eremoleon Banks, 1901

This New World genus contains 17 described species, but probably at least that many more spe-
cies await description. The larvae live under rock overhangs or in cave mouths, usually anchored on
bedrock with fine dust covering them. The species have elongate legs similar to E/achyleon and
Glenurus, but do not have the pretarsal claws modified to fold against the distal tarsomere. Species of
Glenurus have more prominent wing markings and have the posterior gonapophysis of the female
terminalia greatly reduced in size. Two species of Eremoleon found in Costa Rica have the forewing
costal area biareolate, which is uncommon in this genus. It is likely that further species of Eremoleon
will be found in Costa Rica when a search for larvae in caves and rock overhang habitats is done.

284 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES
Volume 53, No. 12

KEY TO COSTA RICAN EREMOLEON

1. Forewing with a prominent arched line originating near hind border and vein Cu (Fig. 636) .......... E. genini
We Rorewingawithoutprominentarched imei (Higs653) eamsueien amen cnn anen nn E. dunklei

Eremoleon dunklei Stange, 1999
(Figs. 632-635)

DIAGNOSIS. — Greatest ocular width about equal to greatest interocular distance (Fig. 634);
forewing without prominent sinuate line (Fig. 633).

GEOGRAPHICAL DISTRIBUTION. — Costa Rica, known only from the holotype collected at
PUNTARENAS: Monteverde.

ALTITUDINAL DISTRIBUTION. — 1400 m.

ADULT FLIGHT PERIOD. — Late February.

HABITAT PREFERENCE. — Unknown.

Eremoleon genini (Navas, 1924)
(Figs. 636-637)

DIAGNOSIS. — Greatest ocular width much less than greatest interocular distance; forewing with
prominent sinuate line (Fig. 636).

GEOGRAPHICAL DISTRIBUTION. — Mexico and Costa Rica: SAN JOSE.

ALTITUDINAL DISTRIBUTION. — Elevation about 1000 m.

ADULT FLIGHT PERIOD. — Unknown.

HABITAT PREFERENCE. — Unknown.

Glenurus Hagen 1866

This is a uniform genus easily recognized in Costa Rica by the considerable brown suffusion of
the hind wing and usually forewing apical field. This color pattern evidently offers camouflage as the
adults fly in and out of the shadows in shaded woods. However, it is apparent that the adults are
noctural as they are attracted to lights, often after midnight. The larvae are unique 1n the New World in
having only two mandibular teeth. They live usually in association with wood, either in dry tree holes
or wood debris in decaying logs. They are slow moving and burrowing except for the faster Glenurus
proi which lives on top of loose, rain-protected soil. The genus has eight described species ranging
from the southeastern United States south to Argentina, with two species recorded from Costa Rica
and a third likely occurring there.

KEY TO COSTA RICAN GLENURUS

Ile Pterothoracic pleura sharply divided into dark brown band above and pale brown band below (book cover) . G. proi
1’. Pterothoracic pleura not divided into dark brown and pale brown bands .......................- 2

2(1'). Hind basitarsus longer than distal tarsomere; forewing without the prominent brown suffusion in the apical field that
isproundmni the thin swine |(RigsO29) ee eee a ewes tenes es eerie ee eer G. heteropteryx

2'. Hind basitarsus shorter than distal tarsomere; forewing with prominent brown suffusion in apical field similar to that
foundanibind wine (ETE 63.0). os sare cee toers oh Seyi aie as as oe eas oss sc G. luniger

PENNY: LACEWINGS OF COSTA RICA 285

Glenurus heteropteryx Gerstaecker, 1885
(Fig. 629)

DIAGNOSIS. — Easily distinguished from other species of G/enurus by the lack of the prominent
brown suffusion in the forewing (Fig. 629). The hind basitarsus is longer than the distal tarsomere,
which is also found in Glenurus proi.

GEOGRAPHICAL DISTRIBUTION. — Described from Panama and Trinidad. Also found in Vene-
zuela and Ecuador. Very likely to be found in Costa Rica.

ALTITUDINAL DISTRIBUTION. — Unknown.

ADULT FLIGHT PERIOD. — Unknown.

HABITAT PREFERENCE. — Unknown.

Glenurus luniger Gerstaecker, 1893
(Fig. 630)

DIAGNOSIS. — The nearly solid brown band from the pterostigma to the hind margin in the
forewing (Fig. 630) is distinctive from G. heteropteryx (lacks the band) and G. proi (has the band bro-
ken up ventrally). The short basitarsus, which is shorter than the distal tarsomere, provides a good
character to separate this species from other Costa Rican species.

GEOGRAPHICAL DISTRIBUTION. — Ranges from southwestern U.S.A. (Arizona and California)
south to Panama.

ALTITUDINAL DISTRIBUTION. — Unknown.

ADULT FLIGHT PERIOD. — Unknown.

HABITAT PREFERENCE. — The larvae of this species have been found living in wood debris asso-
ciated with dry tree holes and also in leaf litter under rock overhangs.

Glenurus proi Navas, 1929
(cover, Fig. 631)

DIAGNOsIS. — The sharp color division of the pterothoracic pleura into a dark brown band above
and pale brown below is distinctive for this species (book cover). The hind basitarsus is longer than the
distal tarsomere, as is also found in G. heteropteryx, which lacks the prominent dark brown stripe in
the apical field. In G. proi, this band is not solid as in G. /uniger and there are pale spots near the ventral
margin of the band.

GEOGRAPHICAL DISTRIBUTION. — Described from Mexico but has also been collected in Costa
Rica: GUANACASTE: Santa Rosa National Park.

ALTITUDINAL DISTRIBUTION. — About 300 m.

ADULT FLIGHT PERIOD. — March and December in Costa Rica.

HABITAT PREFERENCE. — Larvae of this species were found living on top of the soil under a
large, ground-level, tree root overhang in Mexico. Nearby was a rodent burrow, which is significant
since it is believed that ovipositing females are attracted by the odor of animal feces. The larvae are
highly unusual in that they cover themselves with debris, including rodent pellets, and proved to be
relatively active predators compared to other known larvae of Glenurus.

Purenleon Stange, new genus
TYPE SPECIES. — Purenleon abruptus Stange, new species, by present designation.

= Diazus Navas 1914:220 (preoccupied by Diazus Le Conte, 1859, Scarabaeidae: Coleoptera),
by original designation. New synonymy.

286 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES
Volume 53, No. 12

TYPE SPECIES. — Diazus clavatus Navas, by monotypy.

DIAGNOSIS. — Differs from the majority of genera of the tribe Nemoleontini by the stout legs
with the forefemur swollen. Also, the tibial spurs are present, the pretarsal claws are not capable of
closing against the distal tarsomere and there is no postventral lobe developed from the male
ectoproct.

DESCRIPTION. — Antennal fossa separated from ocular rim by much less than the greatest diame-
ter of scape; pronotum wider than long; mesoscutellum smaller than metanotum; legs about equal in
length, hind leg sometimes longer than foreleg; tibial spurs present; basitarsus of hind leg at least 2.5
times longer than greatest width, longer than pretarsal claws; pretarsal claws not capable of closing
against distal tarsomere; forewing with anterior margin evenly curved toward apex, costal area at
point of coalescing of subcostal and radial veins lower than at middle of wing; forewing radial sector
originates well beyond forking of CuA; posterior fork of forewing vein CuA at an oblique angle to
hind margin; forewing vein 2A widely separated from normal 3A before strongly angling toward pos-
terior margin; hind wing vein CuA extends to, or sometimes well beyond, forking of MP; male
ectoproct simple without postventral lobe; male paramere in form of rigid plate; male gonarcus not
greatly expanded laterally; female tergite VIII without well-developed subapical row of long, stout
setae; female ectoproct without digging setae; posterior gonapophysis well developed, often some-
what inflated.

Included species: Purenleon abruptus Stange, P. albovaria (Banks) [new combination], P.
bistictus (Hagen) [new combination], P. clavatus (Navas) [new combination], P. connexus (Banks)
[new combination], P. cubensis (Alayo) [new combination], P. debilis (Gerstaecker) [mew combi-
nation], P. imbellis (Banks) [new combination], P. iniguus (Navas) [new combination], P.
inscriptus (Hagen) [new combination], P. minor (Banks) [new combination], P. nubipennis
(Navas) [new combination], P. parallelus (Banks) [new combination], P. reductus (Banks) [new
combination] and P. zayasi (Alayo) [mew combination].

OBSERVATIONS. — The type species of the genus Psammoleon are now placed in the genus
Euptilon Westwood, a genus misinterpreted by later workers. The holotype of the type species of
Euptilon is apparently missing and the colored figure of the type species shows a composite insect
with antlion wings and body, but with the attached antennae of a corydalid. The green color of the
body is misleading since this color is absent in the Myrmeleontidae. Evidently the insect had some
green mold or perhaps green discoloration of the pin. The wings are clearly those of what has been
called Psammoleon guttipes Banks, which is now considered a synonym of Psammoleon ornatus.
This is especially evident from the type locality of Hemerobius ornatus which is Dinwiddie, Virginia.
No other insect from this area can be confused with Psammoleon ornatus. Psammoleon arizonensis
Banks, P. decipiens Banks, P. ornatus (Drury) (= P. guttipes Banks) and P. sinuatus Currie now con-
stitute the genus Euptilon. Many of the species here included in Purenleon formerly were placed in
Psammoleon. However, the species Psammoleon bipunctatus Navas, Psammoleon cautus (Walker),
and Psammoleon nebulosus (Navas) are not closely related to Euptilon or Purenleon and are here
transferred to the genus Dimarella Banks until their relationships can be more closely studied. They
agree with Dimarella in having a well-developed postventral lobe on the ectoproct and similar male
genitalia, but the head is not modified as in Dimarella. Dimarella appears to be centered in Mexico,
Central America and the Caribbean Islands.

Although records of only two species of Purenleon are known from Costa Rica, there are cer-
tainly other species, especially P. parallelus and P. debilis which are known from Mexico and Pan-
ama. Probably there are other species as well, since no good antlion survey has been made for Costa
Rica. The larvae live in sand. The adults are easily distinguished from other groups in Costa Rica by
the stout legs and the elongate forefemoral sense hair, which is over one-half the length of the femur.
The forefemur is swollen, at its widest point being much wider than interantennal distance, and has
abundant pubescence, especially on the closing face.

PENNY: LACEWINGS OF COSTA RICA 287

KEY TO COSTA RICAN PURENLEON

1. Pretarsal claws of foreleg < one-half length of basitarsus; forewing costal area abruptly broadened near base

PeosS) slorewin pdlenetiiless: thant 2 010) 2) ed seco) Sere) cient ay cicepted cece eu) hil eee oe ce eee P. abruptus
1’. Pretarsal claws of foreleg > length of basitarsus; forewing costal area not abruptly widened near base; forewing
Bate CALETA O LETC ALE CMU saris cs nce ot ees Ge eee) Shoe ATEN esr PA yes So SUS. Ue Gee! Ged gets sspgeens tere 2

2(1'). Vein CuP+1A of forewing running parallel to posterior fork of CuA for a long distance beyond origin of radial

sector; mesofemoral sense hair about one-half length of profemoral sense hair. ............. P. parallelus
2’. Vein CuP+1A of forewing running obliquely to hind margin along posterior fork of CuA at a point near origin of
radial sector (Fig. 639, 640); mesofemoral sense hair as long as profemoral sense hair. ............... 3

3(2'). Hind leg pretarsal claws much longer than basitarsus; hind wing much broader at about 3/4 distance from base than
[syzssealliigy Gor GUA Elihy (LEU E (GF 0) ee pment ecm het tart rae Seager startet lori Mla WO aman n lin Nev are tule P. iniquus
3’. Hind leg pretarsal claws shorter than basitarsus; hind wing not especially broadened toward apex (Fig. 639) P. debilis

Purenleon abruptus Stange, new species
(Figs. 638, 642)

HOLOTYPE. — Male, 37.1 km (23 mi) south Matias Romero, Oaxaca, Mexico, I'V.5.1962, L.
Stange collector (Florida State Collection of Arthropods).

DIAGNOSIS. — This species is distinctive in having the forewing costal area abruptly broadened
near the base. Mid-femoral sense hair about one-half as long as that of forefemur, about one-fourth
length of femur. The relatively small pretarsal claws, which are not more than one-half as long as the
basitarsus of the foreleg, is a diagnostic character in Costa Rica.

DESCRIPTION. — Length of body from vertex to apex of abdomen 17 mm, length of forewing and
hind wing 20 mm.General coloration: Light brown, forewing with dark brown on crossveins where
posterior fork of cubital fork meets posterior margin and at rhegma. Face pale brown with broad dark
brown band above and below antennae, emarginate ventrally at middle; vertex (Fig. 642) mostly grey,
pruinose with nearly black scars; mouthparts pale brown except for dark brown mandibles, stipes and
distal palpimere of labium; antenna brown with pale brown at apices of scape and flagellomeres,
pedicel nearly all brown; pronotum pale brown with extensive dark brown medially, laterally and an-
teriorly; mesoscutum mostly dark brown with pale spot on prescutum, irregularly on each side of
mesoscutellum which is paler posteriorly, small round pale spot laterally near wing base; metascutum
darker with pale brown anteriorly and on posterior margin of metascutellum; thoracic pleura mostly
darker brown dorsally, lighter brown ventrally with ventral area mostly dark brown. Coxae and tro-
chanters mostly pale except small basal darkening on coxae; femora mostly pale on closing face, dark
brown on exterior face; foretibia mostly dark brown on closing face, mostly pale brown on exterior
face with dark brown bands basally, distally and at middle; mid-tibia and hind tibia mostly pale brown
on closing face except apically, mostly dark brown on exterior face. Tarsi dark brown except pale
brown basitarsus which is darker apically. Abdominal tergites mostly dark brown with pale distal
ends, sternites with more pale brown, especially sternites VI to VIII. Ectoproct and sternite LX nearly
all light brown. Wing membranes nearly without suffusion, white stigma and minor dark brown suffu-
sion on crossvein near posterior margin where the posterior branch of cubital fork ends, and at
rhegma; wing veins and crossveins mostly pale brown but with extensive dark brown coloring pro-
ducing mostly an alternating pattern of light brown and dark brown.

CHAETOTAXY. — Clypeus, labrum and postmentum with outstanding, mostly white setae, else-
where on head inconspicuous; setae very short on antennae and mouthparts except distal palpimere of
labium with brown setae about as long as width of distal part of palpimere. Pronotum with erect black
setae, especially submedially and at posterior margin and three to four elongate white setae
sublaterally which are slightly thickened toward apex; rest of nota with few inconspicuous black

288 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES
Volume 53, No. 12

setae; thoracic pleura with numerous elongate, white setae, some of which are slightly thickened to-
ward apex. Forecoxa with many elongate setae posteriorly that are longer than greatest coxal width
and many smaller white setae on lateral surface that are shorter than coxal width; forefemur with many
elongate white setae on closing surface, shorter decumbent ones on exterior surface; several elongate,
erect black setae on closing surface, several subapically on exterior surface; mid-femur with some
white decumbent setae on exterior face and one prominent erect black seta, mesally with many elon-
gate white setae, basal half forming a comb with the setae thickened; hind femur with elongate, erect
black setae on distal one-third of exterior face and on middle one-third of closing face; some short
white setae on most of femur with a few erect ones on closing face; foretibia with many black erect
setae on exterior face and with mostly decumbent white setae, especially on base of exterior face but
with few erect white setae on closing face; mid-femur with scattered elongate, erect black setae, some
of which are longer than mid-femoral sense hair; hind femur with fewest setae, all black, some short
but with many, very long, black setae. Abdomen with predominately short white, semi-decumbent
setae except for ectoproct which has elongate, mostly brown, setae. Wings with short setae, none ex-
ceeding twice the vein diameter.

STRUCTURE. — Head with vertex slightly raised above eyes; greatest ocular width slightly
greater than interocular distance at middle; antenna moderately long and weakly clavate, with 36
flagellomeres, basal flagellomeres about twice as long as wide, others broader than long; distal
palpimere of labium moderately swollen with long narrow apex, palpimacula oval, situated about
midway; pronotum a little wider than long. Legs similar in length, foreleg shorter than midleg which is
shorter than hind leg; forefemur swollen, femoral sense hair long, about half as long as forefemoral
length; foretibia and mid-tibia swollen irregularly, slightly narrower at middle; mid-femoral sense
hair about one-half as long as that of forefemur, about one-fourth length of femur; basitarsus of foreleg
about 3.5 times longer than middle diameter, shorter than distal tarsomere; basitarsus of midleg about
4.0 times longer than middle diameter, and that of hind leg about 6 times longer than middle diameter,
longer than distal tarsomere; tibial spurs of foreleg and midleg reaching to about apex of second
tarsomere, that of hind leg to apex of basitarsus; pretarsal claws short, about as long as tarsomere II of
hind leg. Forewing the same length as hind wing; forewing costal area (Fig. 638) abruptly broadened
at base, costal cells not interconnected by crossveins, those above radial sector about 1.5 times as high
as wide. Abdomen much shorter than forewing length; male ectoproct simple, not produced.

FEMALE. — Similar to the male except for terminalia. Pregenitale well developed; posterior
gonapophysis about four times longer than middle diameter with many elongate setae, some of which
are subequal in length to gonapophysis; lateral gonapophysis without digging setae, usually well con-
cealed in genital complex; ectoproct strongly produced ventrally with row of about four elongate
black setae ventrally, elsewhere many other setae which are finer.

VARIATION. — The length of the forewing varies from about 18 to 25 mm. Forewing rhegmal
area more prominently marked in some specimens.

PARATYPES (16 males, 4 females). — | male, 1 female, Santa Rosa National Park, Guanacaste
Prov., COSTA RICA, 300 m, March 1984, D. H. Janzen and W. Hallwachs (INBIO, Costa Rica); ); 1
male, Rabinal, Baja Verapaz, GUATEMALA, II.7.1965, G. S. O’Neal (FSCA); 1 female, train
64.5 km (40 mi) E Guatemala City, 1V.7.1926, J. M. Aldrich (USNM); 2 males, | female, Sierra de
Agalta, 8 km. NE Catacamas, Olancho, HONDURAS, 3000’, L. Stange and R. Miller (FSCA); same
data as holotype (FSCA; Miller collection; USNM); 1 male, 11.3 km (7 mi) NE Tepanatepec,
Oaxaca, MEXICO, 4.11.1985, L. Stange and R. Miller (FSCA); 3 males, Chamela, Jalisco, MEXICO,
15/24.1V. 1986, F. Parker (FSCA).

BIOLOGY. — Larvae were found living in loose soil under bark and other debris at the base of a
large tree at Matias Romero, Mexico. Four were reared. The duration of pupation varied from 35 to 50
days.

PENNY: LACEWINGS OF COSTA RICA 289

OBSERVATIONS. — This species is very distinctive in having the forewing costal area abruptly
broadened near the base. It belongs to the P. inscriptus group, which is characterized by relatively
small pretarsal claws that are not more than one-half as long as the basitarsus of the foreleg.

GEOGRAPHICAL DISTRIBUTION. — Southern Mexico south to Costa Rica. The only Costa Rican
record is: GUANACASTE: Santa Rosa National Park.

ALTITUDINAL DISTRIBUTION. — 300 m.

ADULT FLIGHT PERIOD. — March.

HABITAT PREFERENCE. — See Biology.

Purenleon debilis (Gerstaecker, 1893) [new combination]
(Figs. 639, 643)

DIAGNOSIS. — This is a medium-sized antlion (body length 20-28 mm; forewing length
28-31 mm) that is closely related to Purenleon iniquus. The pretarsal claws are longer than in that
species and the hind wing (Fig. 639) has a different shape.

GEOGRAPHICAL DISTRIBUTION. — Mexico south to Panama. No Costa Rican records are cur-
rently known, but it will likely be found there.

ALTITUDINAL DISTRIBUTION. — Mid-elevations.

ADULT FLIGHT PERIOD. — August to November in other Central American countries.

HABITAT PREFERENCE. — Unknown.

Purenleon iniquus (Navas, 1914) [mew combination]
(Figs. 640, 644)

DIAGNOSIS. — This species is about the same size as Purenleon debilis and has a similar basic
coloration, although details of the thoracic pattern are paler and more elaborate (Fig. 644). The hind
wing shape (Fig. 640) is different from that species. The pretarsal claws are shorter than in P. debilis.

GEOGRAPHICAL DISTRIBUTION. — Honduras (holotype) south to Panama. Records from Costa
Rica are from Santa Rosa National Park (GUANACASTE) and Caldera.

ALTITUDINAL DISTRIBUTION. — About 400 m.

ADULT FLIGHT PERIOD. — December.

HABITAT PREFERENCE. — The larvae of this species were found living in loose soil around large
trees in Honduras.

Purenleon parallelus (Banks, 1935) [new combination]
(Figs. 641, 645)

DIAGNOSIS. — The forewing venation is distinctive in that the vein CuP+1A runs parallel to the
posterior fork of CuA for a long distance (Fig. 641). This condition is not duplicated in any other de-
scribed Purenleon. The hind wing is shorter than the forewing so that in repose the apices coincide
(hind wing longer than forewing in P. debilis and P. iniquus). Also, the mesofemoral sense hair is only
half as long as that of the profemur, which is another distinguishing character from P. debilis and P.
iniquus. This species is smaller than most other Costa Rican Puren/leon (forewing length 22—24 mm).

GEOGRAPHICAL DISTRIBUTION. — Southern Mexico south to Panama on the Pacific Coast. This
species is not currently recorded from, but probably occurs in Costa Rica and should be looked for in
coastal sand dune localities.

ALTITUDINAL DISTRIBUTION. — Sea level.

ADULT FLIGHT PERIOD. — March to June.

HABITAT PREFERENCE. — The larvae of this species live in sand dune areas on the coast.

290 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES
Volume 53, No. 12

FAMILY POLYSTOECHOTIDAE

by
Norman D. Penny

There are only four extant species of Polystoechotidae known, two species from Chile and two
species from North and Central America. The most widespread species, Polystoechotes punctatus, is
the only species found in Costa Rica.

Polystoechotes punctatus (Fabricius, 1787)
(Fig. 658)

DIAGNOSIS. — This species is a most distinctive neuropteran, being large and broad-winged,
with many radial sectors in the forewing (Fig. 658). It does not have the long antennae of ascalaphids,
and wings are much broader than antlions.

GEOGRAPHICAL DISTRIBUTION. — The geographical distribution of this species historically in-
cluded almost all of cool, temperate North America and the high mountain chains of Central America
and Mexico as far south as Panama (Penny, unpublished data). In the past half century it appears to
have disappeared from much of its former range in temperate North America, including all of the east-
ern U.S.A. and Canada. Within Costa Rica it has only been collected from Cartago (CARTAGO) and
Monteverde (PUNTARENAS).

ALTITUDINAL DISTRIBUTION. — 1500 m at Monteverde.

ADULT FLIGHT PERIOD. — Mid-May to July.

HABITAT PREFERENCE. — In Central America this species appears to be associated with
montane forest.

FAMILY SISYRIDAE

by
Norman D. Penny

Spongilla-flies are rather small neuropterans with aquatic larvae that feed on freshwater sponges.
Adults usually have a short emergence period. In the central Amazon Region, adult emergence is usu-
ally timed to the seasonal rise and fall of rivers and streams, with peak emergence occurring at, or
shortly after, the high water period (July). Many species are attracted to lights and can be collected
some distance from the nearest stream. Adults superficially resemble brown lacewings
(hemerobiids), but have only a single radial sector. The single species presently known from Costa
Rica is Sisyra apicalis, although others probably occur there.

Sisyra apicalis Banks, 1908
(Figs. 659, 660)

GEOGRAPHICAL DISTRIBUTION. — Originally described from Cuba this species 1s known from
U.S.A. (Gulf Coast) as far south as Panama (Parfin and Gurney 1956) andthe Brazilian Amazon
(Penny 1981). Within Costa Rica only one collection record is known from: GUANACASTE: 14 km
S Canas.

ALTITUDINAL DISTRIBUTION. — Below 200 m near Canas.

ADULT FLIGHT PERIOD. — February.

PENNY: LACEWINGS OF COSTA RICA 291

HABITAT PREFERENCE. — Near Cafias this species has been captured at lights near a me-
dium-sized river with patchy forest and open fields surrounding it.

292

FIGURES

A list of abbreviations used with the figures is as follows:

1A = first anal vein
2A =second anal vein
3A = third anal vein

a = aedeagus

acu = acumen

arc = arcessus

ax = axillary angle

C =costa

ce = callus cerci

cf = costal field

CuA = anterior cubitus
CuC = cubital cell
CuP = posterior cubitus
d = discal area

dapo = dorsal apodeme
dp = distal process

ect = ectoproct

ent = entoprocessus

f = frenulum

gc = gonarcus

gcr = gonocristae

gcu = gonocornua

gdh = gonarcus dorsal horns

gps = gonapsis

gsc = gonosaccus

gst = gonosetae

h = humeral crossvein
hy = hypandrium

hyt = hypandrium transverse plate

ig = inner gradate series

jl = jugal lobe

la = lateral arm of gonarcus
Ip = labial palpus

MA = anterior media

PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES

Volume 53, No. 12

mai = median apical incision of hypandrium

mi = microtholi

mp = maxillary palpus
MP = posterior media
mu = mediuncus

og = outer gradate series
p = penis

pa = parameres

ps = pseudopenis

pt = processes terminales
r-m = radial-medial crossveins
r-r = radial crossveins

Ry, = radius
Rs = radial sector
s = seta

Ss +9 = eighth + ninth sternites
So = ninth sternite

sap = subanal plate

Sc = subcosta

scf = subcostal field

st = pterostigma

sty = stylus

To9+ect = 9th tergite + ectoproct
ti = tignum

vapo = ventral apodeme

PENNY: LACEWINGS OF COSTA RICA 293

Ficure |. Ululodes macleayana in habitus view. (Illustration by Artémio Coelho da Silva)

FicurE 2. Neohaploglenius flavicornis, lateral view of head. (Illustration by Victoria Saxe)
FiGuRE 3. Ululodes macleayana, lateral view of head. (Illustration by Victoria Saxe)

294 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES
Volume 53, No. 12

repagula

Figure 4. Ululodes sp. eggs and larvae. (Illustration by Artémio Coelho da Silva)

PENNY: LACEWINGS OF COSTA RICA 295

)

‘
a
HS oe oe}

FiGure 5. Undetermined Haplogleniinae larva. (Illustration by Alberto Coelho da Silva)

296 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES
Volume 53, No. 12

ean

Se,

LE Ss SES Ds Te

sss

Hy wat
ITTY TLE sereese sate $3) ses .
eS.
pees See eee
AIS oe oH
LA os ASR Sa Os F

FicurE 6-11. Right fore- and hindwings of Haplogleniinae. Fig. 6, Amoea vacuus; Fig. 7, Haploglenius luteus; Fig. 8,
Ascalobyas albistigma; Fig. 9, Haploglenius peruvianus; Fig. 10, Ascalobyas microcerus, Fig. 11, Neohaploglenius
flavicornis. (Mlustrations by Linda Seabrooks Campbell)

PENNY: LACEWINGS OF COSTA RICA

Se nd IES ES SS Sea
=e

See

pe a
SS aNpgare

297

or Pere
Prec hd
“ayauecaeeaagees

Sees

IS
preweres saeseees

(13-16)]

Figures 12-16. Right fore- and hindwings of Ameropterus. Fig. 12, A. breviantennis; Fig. 13, A. consors; Fig. 14, A.

mexicanus; Fig. 15, A. subripiens; Fig. 16, A. trivialis. [Mlustrations by Jere Schweikert (12) and Linda Seabrooks Campbell

‘

298 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES

4
4
Volume 53, No. 12 |

vrei ieee
—SstaF
SY
WOT

Ta Mie a a a

a eee eee ee

Ficures 17-21. Right fore- and hindwings of Cordulecerus. Fig. 17, C. inquinatus (male); Fig. 18, C. inquinatus (female);
Fig. 19, C. praecellens (male); Fig. 20, C. praecellens (female); Fig. 21, C. subiratus. [Illustrations by Linda Seabrooks Camp-
bell (17—20) and Jere Schweikert (21)]

PENNY: LACEWINGS OF COSTA RICA 299

= a Be asig we
Fe ae eS a ae as rg.
peseastesge
ERE EES

z
aS

Tas ‘3 ae aeeeeeeese"
Sane eases Co Te ig
20 Nests og CUR

ee Sat SB is so en CPE IS TP RE
WR TR

ee
reueees SEES See

FiGuRES 22-26. Right fore- and hind wings of Ululodes. Fig. 22, U. bicolor (male); Fig. 23, U. bicolor (female); Fig. 24, U.
cajennensis; Fig. 25, U. macleayana (male); Fig. 26, U. macleayana (female). (Illustrations by Linda Seabrooks Campbell)

300 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES
Volume 53, No. 12

Mee Ge biggie

ata aes

(hig AILS

OEE Lie

eo
ntads\ eae
Ait Ain b ae,
TV tifa das eee
FAAS

FIGURE 27. Spiroberotha sanctarosae, right fore- and hind wings. (Illustration by Jere Schweikert)

PENNY: LACEWINGS OF COSTA RICA

—_ |
PA Sc R1
‘ eh DS
RY RES!

| PALD . ™ #; ae a
| lone piv OG ae eee
LULA TESS

VQ
\ A ~ — eS .

FIGURE 28. Ceraeochrysa elegans, habitus view. (Illustration by Victoria Saxe and Virginia Kirsch)

302 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES
Volume 53, No. 12

FiGuRE 29. Generalized schematic view of a male chrysopid genitalia. 9S = ninth sternite; acu = acumen; ap-ect = dorsal
apodeme of ectoproct; ent = entoprocessus; gcr = gonocristae; gp = gonapsis; gs = gonarcus; gsac = gonosaccus; la-gs = lateral
arm of gonarcus; mi=microtholi; mu=mediuncus; ps = pseudopenis; sap = subanal plate; ti= tignum. (from Adams and Penny,
1987)

PENNY: LACEWINGS OF COSTA RICA

SSG
ATT

clypeus
labrum

FIGURES 30-32. Domenechus mirifica (Gerstaecker). Fig. 30, right fore- and hind wings; Fig. 31, head in frontal view; Fig.
32, head and thorax in dorsal view. (Illustrations by Virginia Kirsch)

304 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES
Volume 53, No. 12

”
7/

SSS

35 ,

FiGuRES 33-35. Loyola croesus (Gerstaecker). Fig. 33, right fore- and hind wings; Fig. 34, head in frontal view; Fig. 35, head
and thorax in dorsal view. [Illustrations by Fred Ehrman (33, 35) and Diane Wong (34)]

PENNY:

LACEWINGS OF COSTA RICA

nN
Ss

we
Sj

a
<a
BAA) SS

“E>
Saeser Quakes
ipa ee
RGSS

Thess

FiGuREs 36-38. Berchmansus elegans (Guérin-Ménéville). Fig. 36, habitus dorsal view; Fig. 37, head in frontal view; Fig.
38, head and thorax in dorsal view. [Illustrations by Virginia Kirsch (36) and Fred Ehrman (37, 38)]

305

306 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES
Volume 53, No. 12

FIGURES 39-41. Gonzaga palliceps (McLachlan). Fig. 39, right fore- and hind wings; Fig. 40, head in frontal view: Fig. 41,
head and thorax in dorsal view. (Illustrations by Fred Ehrman)

PENNY: LACEWINGS OF COSTA RICA 307

ksh ="

42 BESS

FiGuRES 42-44. Gonzaga torquatus Navas. Fig. 42, right fore- and hind wings; Fig. 43, head in frontal view; Fig. 44, head
and thorax in dorsal view. (Illustrations by Fred Ehrman)

308 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES
Volume 53, No. 12

ETE EEE ee

TTT TLL
ROSES Ve
SSS

FIGURES 45-47. Leucochrysa (Leucochrysa) adamsi Penny. Fig. 45, right fore- and hind wings; Fig. 46, head in frontal view;
Fig. 47, head and thorax in dorsal view. [Illustrations by Virginia Kirsch (45) and Norman Penny (46, 47)]

PENNY: LACEWINGS OF COSTA RICA 309

a a
SL
Sil

URS

FIGURES 48-50. Leucochrysa (Leucochrysa) clara (McLachlan). Fig. 48, right fore- and hind wings; Fig. 49, head in frontal
view; Fig. 50, head and thorax in dorsal view. (Illu i d Ehrman

a 48,
al v (Illustrations by Fred Ehrman)

310 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES
Volume 53, No. 12

FiGuRES 51—53. Leucochrysa (Leucochrysa) colombia (Banks). Fig. 51, right fore- and hind wings; Fig. 52, head in frontal
view; Fig. 53, head and thorax in dorsal view. (Illustrations by Virginia Kirsch)

PENNY: LACEWINGS OF COSTA RICA 311

TTT

FiGuRES 54—56. Leucochrysa (Leucochrysa) lestagei Navas. Fig. 54, right fore- and hind wings; Fig. 55, head in frontal view;
Fig. 56, head and thorax in dorsal view. (Illustrations by Fred Ehrman).

312 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES
Volume 53, No. 12

099

FIGURES 57-59. Leucochrysa (Leucochrysa) magnifica (Banks). Fig. 57, right fore- and hind wings; Fig. 58, head in frontal
view; Fig. 59, head and thorax in dorsal view. (Illustrations by Fred Ehrman)

PENNY: LACEWINGS OF COSTA RICA 313

Es
Bie

FIGURES 60-62. Leucochrysa (Leucochrysa) pretiosa (Banks). Fig. 60, right fore- and hind wings; Fig. 61, head in frontal
view; Fig. 62, head and thorax in dorsal view. (Illustrations by Fred Ehrman)

314 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES
Volume 53, No. 12

VER

Rare ae ate meneee: (te
RRS
Seeks
63 XU = |

FIGURES 63-65. Leucochrysa (Nodita) amazonica Navas. Fig. 63, right fore- and hind wings; Fig. 64, head in frontal view;
Fig. 65, head and thorax in dorsal view. [Illustrations by Diane T. Sands (63) and Zena Deretsky (64, 65)]

PENNY: LACEWINGS OF COSTA RICA 315

FIGURES 66-68. Leucochrysa (Nodita) amistadensis Penny. Fig. 66, right fore- and hind wings; Fig. 67, head in frontal view;
Fig. 68, head and thorax in dorsal view. [Reprinted from Penny (2001)]

316

FiGuRES 69-71. Leucochrysa (Nodita) antica Navas. Fig. 69, right fore- and hind wings; Fig. 70, head in frontal view; Fig.
71, head and thorax in dorsal view. [Illustrations by Renata Zakhvatkina (69) and Virginia Kirsch (70, 71)]

PENNY: LACEWINGS OF COSTA RICA 317

eae
aN it

‘a

Cee ee

FiGURES 72—74. Leucochrysa (Nodita) askanes (Banks). Fig. 72, right fore- and hind wings; Fig. 73, head in frontal view;
Fig. 74, head and thorax in dorsal view. [Illustrations by Diane T. Sands (72) and Victoria Saxe (73, 74)]

318 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES )
Volume 53, No. 12 |

ee ee a

FIGURES 75-77. Leucochrysa (Nodita) caucella Banks. Fig. 75, right fore- and hind wings; Fig. 76, head in frontal view; Fig.
77, head and thorax in dorsal view. [Illustrations by Sarah Donelson (75, forewing), Renata Zakhvatkina (75, hind wing), and
Virginia Kirsch (76, 77)]

PENNY: LACEWINGS OF COSTA RICA 319

Pars,

EE
id

FIGURES 78-80. Leucochrysa (Nodita) compar (Navas). Fig. 78, right fore- and hind wings; Fig. 79, head in frontal view; Fig.
80, head and thorax in dorsal view. [Illustrations by Virginia Kirsch (78) and Victoria Saxe (79, 80)]

320 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES
Volume 53, No. 12

FIGURES 81-83. Leucochrysa (Nodita) cornesta (Banks). Fig. 81, right fore- and hind wings; Fig. 82, head in frontal view;
Fig. 83, head and thorax in dorsal view. [Illustrations by Diane T. Sands (81) and Zena Deretsky (82, 83)]

|
|
’

PENNY
: LACEW
INGS OF COSTA RICA
321

TELLERS
Te

. Leucoch
: a) duarte B = g
dc yrSal VIEW llustrati y rl :
Q 5 ig. 85
ew; 1g.

322 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES _
Volume 53, No. 12 +

PEE EEE

FIGURES 87-89. Leucochrysa (Nodita) egregia Navas. Fig. 87, right fore- and hind wings; Fig. 88, head in frontal view; Fig.
89, head and thorax in dorsal view. [Illustrations by Virginia Kirsch (87) and Zena Deretsky (88, 89)]

PENNY: LACEWINGS OF COSTA RICA

, yaa a
a

UE nn
a lay

Ficures 90-92. Leucochrysa (Nodita) forcipata Penny. Fig. 90, right fore- and hind wings; Fig. 91, head in frontal view; Fig.

92, head and thorax in dorsal view. [Illustrations by Virginia Kirsch (90) and Victoria Saxe (91, 92)].

323

324 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES
Volume 53, No. 12 -

55 oe a

Ficures 93-95. Leucochrysa (Nodita) indiga Navas. Fig. 93, right fore- and hind wings; Fig. 94, head in frontal view; Fig.
95, head and thorax in dorsal view. [Reprinted from Penny (2001)]

PENNY: LACEWINGS OF COSTA RICA 325

FiGuREs 96-98. Leucochrysa (Nodita) laertes (Banks). Fig. 96, right fore- and hind wings; Fig. 97, head in frontal view; Fig.
98, head and thorax in dorsal view. [Illustrations by Virginia Kirsch (96) and Victoria Saxe (97, 98)]

326 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES
Volume 53, No. 12

EE
BEET TESS
SATIS”

yas

ae
SS

GGsi

FiGURES 99-101. Leucochrysa (Nodita) lancala (Banks). Fig. 99, right fore- and hind wings; Fig. 100, head in frontal view;
Fig. 100a, head in lateral view; Fig. 101, head and thorax in dorsal view; Fig. 101a, abdomen in dorsal view. [Illustrations by
Virginia Kirsch (99) and Sergio da Freitas (100, 100a, 101, 101a)]

PENNY: LACEWINGS OF COSTA RICA 327

pastieaset

104

Ficures 102-104. Leucochrysa (Nodita) lenora (Banks). Fig. 102, right fore- and hind wings; Fig. 103, head in frontal view;
Fig. 104, head and thorax in dorsal view. [Illustrations by Virginia Kirsch (102) and Michelle Schwengel (103, 104)]

em)
ae

FiGures 105-107. Leucochrysa (Nodita) navasi Kimmins. Fig. 105, right fore- and hind wings; Fig. 106, head in frontal
view; Fig. 107, head and thorax in dorsal view. [Illustrations by Victoria Saxe (105, forewing; 106, 107) and Virginia Kirsch
(105, hind wing)]

PENNY: LACEWINGS OF COSTA RICA 329

etre LL ee

ages

FiGuRES 108-110. Leucochrysa (Nodita) nigrovaria (Walker). Fig. 108, right fore- and hind wings; Fig. 109, head in frontal
view; Fig. 110, head and thorax in dorsal view. [Modified from Penny (2001)}

330 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES
Volume 53, No. 12

FiGures 111-113. Leucochrysa (Nodita) oenops Adams. Fig. 111, right fore- and hind wings; Fig. 112, head in frontal view;
Fig. 113, head and thorax in dorsal view. [Illustrations by Virginia Kirsch (111) and Norman Penny (112, 113)]

PENNY: LACEWINGS OF COSTA RICA a5

FL,

Ficures 114-116. Leucochrysa (Nodita) pallescens (Banks). Fig. 114, right fore- and hind wings; Fig. 115, head in frontal
view; Fig. 116, head and thorax in dorsal view. [Illustrations by Virginia Kirsch (114) and Zena Deretsky (115, 116)]

|
|
332 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES
Volume 53, No. 12
|

|
FIGURES 117-119. Leucochrysa (Nodita) postica Navas. Fig. 117, right fore- and hind wings; Fig. 118, head in frontal view;
Fig. 119, head and thorax in dorsal view. [Illustrations by Diane T. Sands (117) and Victoria Saxe (118, 119)]

PENNY: LACEWINGS OF COSTA RICA 333

Ficures 120-122. Leucochrysa (Nodita) ratcliffei Penny. Fig. 120, right fore- and hind wings; Fig. 121, head in frontal view;
Fig. 122, head and thorax in dorsal view. [Reprinted from Penny (2001)]

334 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES
Volume 53, No. 12

eet

FiGureEs 123-125. Leucochrysa (Nodita) serrei (Navas) (sp. E). Fig. 123, right fore- and hind wings; Fig. 124, head in frontal
view; Fig. 125, head and thorax in dorsal view. [Illustrations by Virginia Kirsch (123) and Norman Penny (124, 125)]

PENNY: LACEWINGS OF COSTA RICA 335

FiGureEs 126-128. Leucochrysa (Nodita) virginiae Penny. Fig. 126, right fore- and hind wings; Fig. 127, head in frontal
view; Fig. 128, head and thorax in dorsal view. [Reprinted from Penny (1998)]

336 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES
Volume 53, No. 12

131

FIGURES 129-131. Vieira leschnaulti (Navas). Fig. 129, right fore-

Internet Archive Audio

Featured

Top

Images

Featured

Top

Software

Featured

Top

Books

Featured

Top

Video

Featured

Top

Mobile Apps

Browser Extensions

Archive-It Subscription

Save Page Now

Full text of "Proceedings of the California Academy of Sciences, 4th series"

See other formats